Skip to main content

Full text of "Rhodora"

See other formats




Conducted and published for the Club, by 


f Associate Editors 

Vol. 76 March, 1974 No. 805 

The Phytogeography and Systematics of Xanthisma tex- 

anum D. C. (Asteraceae); Proper Usage of Intraspecific 

John C. Semple EMEND 1 
A White-Flowered Form of Utricularia purpurea 

C. Barre Hellquist eet 19 
On the Scientific Name of the Longleaf Pine 

BEE Ey Ward T R in 20 
The Occurrence of Bald Cypress (Taxodium distichum (L.) 

Richard) in Suffolk County, Long Island, New York 

Edward J. Hehre and Peter M. Bolid Leeds: 25 

The Nef England Botanical Club, Ine. 
Botanical Museum. Oxford St.. Cambridge, Mass. 02188 

RHODORA. — A quarterly journal of botany, devoted primarily to the 
flora of North America and floristically related areas. Price $10.00 
per year, net, postpaid, in funds payable at par in the United States 
currency at Boston. Some back volumes, and single copies are 
available. For information and prices write RHODORA at address 
given below. 

Scientific papers and notes, relating directly or indirectly to the 
plants of North America, will be considered by the editorial com- 
mittee for publication. Articles concerned with systematic botany 
and cytotaxonomy in their broader implications are equally accept- 
able. All manuscripts should be double-spaced throughout. Please 
conform to the style of recent issues of the journal. Illustrations 
can be used only if the cost of engraver's blocks is met through the 
author or his institution. Forms may be closed five weeks in advance 
of publication. Extracted reprints, if ordered in advance, will be 
furnished at cost. 

Address manuscripts and proofs to Albion R. Hodgdon, 

Dept. of Botany, Nesmith Hall, University of New Hampshire, 
Durham, New Hampshire 03824 

Subscriptions and orders for back issues (making all remittances 

payable to RHODORA) should be sent to RHODORA, Botanical 

Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive 

the next number of RHODORA, changes of address must be received 

prior to the first day of March, June, September or December. 
Second Class Postage Paid at Boston, Mass. 




Vol. 76 March, 1974 No. 805 



The monotypic genus Xanthisma DC. (Asteraceae) in- 
cludes sufficient variation in its single species X. texanum 
DC. to illustrate the three levels of taxonomically recog- 
nized infraspecific variation: subspecifie, varietal and for- 
mal. Proper usage of the three taxonomic ranks requires 
consideration of morphological and distributiona] data ac- 
cording to a consensus of international opinion. Although 
much has been published on the B-chromosome of the spe- 
cies (see Semple, 1972a), no modern systematic treatment 
of X. texanum has been published. Examination of some 
five hundred herbarium collections and experimental data 
indicate that a revision of Gray's 1852 treatment, which 
was based on only a few specimens, is necessary. 

A detailed diseussion of the distribution of the species 
as a whole and each of the infraspecific taxa is warranted 
for two reasons: first, to document the allopatric condition 
of the two subspecies, and second, to show that the distri- 

"The work was part of an NDEA Title IV Fellowship sponsored 
doctoral dissertation submitted to Washington University, St. Louis, 
Mo. Aug. 1972: Dr. Walter H. Lewis advisor. 

2Current Address: Dept. of Biology, University of Missouri-St. 
Louis, St. Louis, Mo. 63121. 

2 Rhodora [Vol. 76 

bution of the species is not predictable from the general- 
ized vegetation distributions given by Gould (1962). The 
Edwards Plateau and its eroded portions are basic con- 
siderations in both cases. A taxonomic treatment consider- 
ing all aspects of the problem is given at the end of the 
discussion of distribution and morphological variation. 

All populations of a species ought to be interfertile to 
some degree. The validity of treating all the morphological 
forms of Xanthisma as a single species was tested by hy- 
bridization experiments involving greenhouse plants grown 
from fruiting material collected at wild populations. No 
barriers to fertilization were found and normal chromosome 
homologue pairing in meiotic prophase was observed in F, 
generation hybrids, which also set viable seed. Cross pol- 
lination was accomplished following the procedure of Smith 
& Parker (1971) and was facilitated by self-incompatibil- 
ity. Selfing could be induced, but yielded progeny of low 
fitness. Clearly, Gray's combining all taxa of Xanthisma 
into a single species is acceptable. 


Xanthisma texanum grows in a portion of the central 
southwest of the United States: Texas, Oklahoma and east- 
ern New Mexico. The known distribution of the species is 
illustrated in Figure 1 and is based on herbarium collections 
from GH, KSU, MO, NY, OKLA, SMU, TEX, TTC, US, WIS, and 
Arizona State University-Tempe.’ Collections of subsp. 
drummondii (T. & G.) Semple and subsp. texanum are rep- 
resented by dots and stars respectively; var. texanum and 
var. orientalis Semple of subsp. texanum are represented 
by encircled and unenclosed solid stars; putative hybrids 
between the two subspecies are represented by open stars. 
Treatment of Gray's (1852) varieties as subspecies, a term 
he did not use (Fernald, 1940), is based upon involucral 
bract morphology and the nearly allopatric distribution of 
the two taxa, while the distribution and bract morphology 

"The assistance of Frederick Utech and Timothy Love at the last 
two herbaria, respectively, is gratefully acknowledged. 

1974] Xanthisma — Semple 3 

Distribution of 
Xanthisma texanum DC 
ye subspeci(ic hybrids 
subsp drummondii 
@® subsp texanum var texanum 

* subsp texanum var. orientalis 


Figure 1. The distribution of Xanthisma texanum DC. Locations 
of collections have been determined from herbarium collections from 
eleven herbaria cited in the text. A key to the symbols used is given. 
Collections are plotted on a county outline map of Texas and Okla- 

4 Rhodora [Vol. 76 

variation within subsp. texanum is properly treated at the 
varietal level. Variation in environmental factors accounts 
for the species distribution. 

Climatic factors determine the extent of the species’ 
range in the east, northwest and south. Too much rain in 
the east and too little in the north and west are the ap- 
parent limiting factors, while increasingly tropical condi- 
tions become limiting to the south (Visher, 1954). Local 
variation in geological factors accounts for the distribution 
within the range and secondarily along the margins of the 
range. The western margin of the eastern deciduous forest 
(Bray, 1906; Tharp, 1926; Braun, 1950; and Gould, 1962) 
forms the eastern margin of the range of X. texanum, 
which is a member of the ecotone flora sensu latu between 
the forests and grasslands. Reasons for the present loca- 
tion of the forest-prairie ecotone are not appropriate for 
discussion in this paper. 

A detailed treatment of geological influences on the spe- 
cies’ range is given by Semple (1972b) and summarized 
for each taxon under each appropriate heading. The Rio 
Grande Valley supports a noticeably distinct vegetation 
community (Foscue, 1932). Xanthisma texanum is excluded 
from the community, which apparently forms a barrier to 
southward expansion of the range of X. texanum, since no 
collections have yet been made in Mexico. Either the Rio 
Grand Valley community is an impassible barrier or no 
inhabitable locations exist in Mexico for X. texanum. 


A subspecies is characterized by all members exhibiting 
a particular morphology distinct from other individuals in 
the species and by the allopatric distribution of these mem- 
bers from the rest of the species (Du Reiz, 1930; Clausen, 
1941; van Steenis, 1957; Valentine & Lóve, 1958; and 
Ehrendorfer, 1968). Those morphologically intermediate 
individuals between the subspecies and the rest of the spe- 

1974] Xanthisma — Semple 5 

$ € 

Figure 2. Geographie distribution of involucral bract variation of 
Xanthisma texanum DC. Bracts from plants from 68 populations are 
illustrated in outline on a map of Texas and Oklahoma. Bracts 1-48 
are from individuals of subsp. drummondii (T. & G.) Semple; bracts 
52-68 are from individuals of subsp. texanwm; and bracts 49-51 are 
from putative hybrids between the two subspecies. 

6 Rhodora [Vol. 76 

cies only occur in populations geographically located be- 
tween the subspecies and the remainder of the species. 
Within X. texanum all members of subsp. drummondit 
(T. & G.) Semple are distinguished by involucral bracts 
that are lanceolate and taper to an acute apex. Subspecies 
texanum includes other members of the species and is char- 
acterized by involucral bracts that are depressed-ovate in 
shape and shorter than those of subsp. drummondii. 
Diagrammatic representations (Figure 2) of involucral 
bracts typical of those occurring in the middle of the series 
forming the involucre are superimposed on an outline map 
of Texas and Oklahoma and were taken from plants occur- 
ring in wild populations at the locations indicated. Bracts 
1-48 show the similarity that occurs within subsp. drum- 
mondii (see Figure 3.C for a detailed illustration of a typi- 
cal involucral bract). Bracts 52-68 illustrate the range of 
variation in involucral bract shape found in subsp. texanum 
(Figures 3.A and 3.B), Bracts 49-51 illustrate involucral 
bracts from putative hybrids between the two subspecies 
and are the only bracts that are intermediate in shape. 
The allopatric distribution of the two subspecies is due 
to the presence of the Edwards Plateau and the Blacklands 
Prairies between the two taxa. Geologically and topo- 
graphically the Edwards Plateau is broken up into five 
regions (Fenneman, 1931; Oetking, 1959; and Raisz, 1957), 
although Gould (1962) included all the varied vegetation 
types within a single vegetation region. With a few excep- 
tions, in part due to man's influences, subsp. drummondii 
does not grow in areas overlying uneroded portions of the 
Edwards Plateau, since the upper strata limit the amount 
of water available to the shallow rooted plants. Depending 
upon the extent of erosion and the nature of the underlying 
bedrock, subsp. drwmmondii grows in varying degrees of 
commonness in the areas of the eastern portion of the 
plateau (colloquial sense) in the following manner: com- 
mon at the lower elevations of the Central Mining District, 
common throughout the Lampasas Cut Plains and rare on 
the more northeastern Comanche Plateau. Both subspecies 

1974] Xanthisma — Semple 7 

are absent in the southeastern portion of the Edwards 
Plateau (the Texas Hill Country). Thus differences in ge- 
ology account for the different numbers of collections in this 
well botanized area and also explain the means by which 
the two subspecies are isolated north and west of San 
Antonio, Texas. 

The presence of the Blacklands Prairies isolates the two 
subspecies north and east of San Antonio. The prairies, 
which run from San Antonio to Dallas and further north- 
eastward, are characterized by dark, high clay content soils 
(Sellards et al., 1966) and are dry and rock hard in the 
summer. Typically, Xanthisma occurs on sandy, light 
brown or reddish soils. Field work in the central Texas 
region showed that X. texanum is very rare throughout 
the entire length of the prairies, except for a few individu- 
als surviving on disturbed or introduced soils along high- 
ways and river banks. Clearly, Gray's varieties are exem- 
plary of what are today treated as subspecies. 

A narrow zone of isolated subsp. drummondii populations 
occurs in Caldwell, Gonzales and Wilson Counties, but the 
small populations only thrive in favorably moist years. 
Interestingly, Heterotheca pilosa (Nutt.) Shinners and 
Bradburia hirtella T. & G. are common in these counties 
and further east, the former being the most prevalent yel- 
low flowered member of the Astereae. Similarity in super- 
ficial appearance of these two species with X. texanum 
undoubtedly explains confusion about the distribution of 
X. texanum in this critical region between the two sub- 


A variety is characterized by all members of a population 
exhibiting a particular morphology distinct from other in- 
dividuals in the species. The distribution of these popula- 
tions is sympatric with populations whose members are not 
within the same variety, and also many populations of 

8 Rhodora [Vol. 76 

morphological intermediates exist (Du Reiz, 1930; Clausen, 
1941; van Steenis, 1957; Valentine & Lóve, 1958; and 
Ehrendorfer, 1968). By this definition varieties can be 
recognized within a species, which does not have subspecies. 
Furthermore, variation that is mot known to be confined 
to an isolated group of populations can not be given sub- 
specific status. Van Steenis (loc. cit.) described varietal 
level variation as being continuous with other varieties, 
although the continuum would have pronounced modes. 
The variation between subspecies can be described as dis- 
continuous, except for the few hybrids. Figure 1 illustrates 
the sympatric condition of the two varieties of subsp. 
texanum: var. texanum with obtuse involucral bract apices 
(Figure 3.A) and var. orientale Semple with cuspidate 
involucral bract apices (Figure 3.B). 


Formal level morphological variation is generally re- 
garded as some phenotype possessed by only some members 
of a population, whereas the subspecific and varietal level 
phenotypes are possessed by all members of a population. 
Those individuals in populations of subsp. texanum var. 
orientale which exhibit red pigmentation along the lateral 
margins of the involucral bracts forming the middle and 
inner series of the involucre have been designated as forma 
rubrum Semple (Figure 3.B). Such individuals are rare, 
but do occur in populations throughout the range of var. 


Experiments were conducted to show that all diagnostic 
characters have a genetic basis. Size but not general shape 
of the involucral bracts was found to be variable. Lack of 
sufficient light, available water, amount of soil or a high 
degree of crowding all resulted in smaller bracts being 
formed than those bracts formed by the individual when 
sufficient quantities of soil, light and water were provided. 

1974] Xanthisma — Semple 9 

The maximum size was determined by heredity and not the 
environment. Therefore size should be considered second- 
arily to shape in aetermining the proper identity of a par- 
ticular collection. 

The red pigment of f. rubrum was found to be passed 
from one generation to the next in var. orientale, but was 
never seen in var. texanum or in subsp. drummondii. Al- 
though the pigment is not always present in detectable 
quantities, its importance warrants some taxonomic status, 
since Xanthisma has always been previously described as 
monochromous. However, in some experimental plants 
heads were occasionally produced which manifested the 
red pigment in the distal portion of the ray straps on the 
adaxial surface and on the lobes of the disc florets, but only 
on the last day or two of disc anthesis. Most individuals 
produced enough pigment to give the rays an orange hue, 
but one plant produced rays that were deep rust red. Such 
variation in intensity of color does not occur on bract mar- 
£ins. No documentation of bicolor rays in wild plants 


Despite the great range of variation in habit occurring 
in the species, no one form was found to warrant taxonomic 
recognition. Within a single population long-stemmed and 
short-stemmed individuals were found in nature. The dis- 
tribution patterns of the short-stemmed plants (Figure 
4.C) and the stemless plants (Figure 4.D) suggest that 
these forms are ecotypes, since they have a genetic basis. 
The extremes of short-stemmedness and stemlessness only 
occur in populations of var. orientale and western popula- 
tions of subsp. drummondii, respectively. The existence of 
à full range of intermediates between the extremes and the 
long-stemmed forms (Figures 4.B, 4.F and 4.G) in most 
populations makes taxonomic recognition unjustified. 

[Vol. 76 




npuowwnjp "esans 

Sijejuaio “ea WAVEIA) "fSqns 

whuera) “eA = whueray ULL 

1974] Xanthisma — Semple 11 


The taxonomic treatment that follows is based on mor- 
phology and distribution. Two dimensional paper chroma- 
tograms were run on leaf and flower extracts of specimens 
of each taxon. In general, subspecies texanum had fewer 
compounds than subspecies drummondii, and var. orientale 
has fewer compounds than var. texanum. In var. orientale 
individuals of forma rubrum had one compound, an uniden- 
tified anthocyanin (Semple, 1972b), not found in other 
individuals of the species. The full significance of the 
chemical studies can only be known upon complete identifi- 
cation of each compound. The preliminary results show 
that chemical data support the taxonomy. The chemical 
studies will have meaning in work on determining the evo- 
lution of the taxa described. 


The most recently published treatment of the genus Xan- 
thisma is in Correll & Johnston (1970). The following 
detailed description of the genus is more complete and ac- 

Figure 3. Involucral Bracts of Infraspecific Taxa. Each bract 
shown. is typical of those in the middle of the series forming the 
involucre: (A) subsp. texanum var. texanum; (B) subsp. texanum 
var. orientale Semple (dashed lines indicate the extent of red pigment 
on the margins of bracts of f. rubrum Semple); (C) subsp. drum- 
mondii (T. & G.) Semple. 

Figure 4. Habit Variation of Xanthisma texanum DC. The base- 
line under each silhouette equals one decimeter. Each form shown il- 
lustrates the habits of experimental plants and represents naturally 
occurring forms. Plants were grown under similar conditions from 
fruiting material collected at wild populations. All plants were from 
different populations, except (B) and (C). See the text for a full 

Figure 5. Heterocarpic Fruit of Xanthisma texanum DC. Ray and 
disc florets produce different kinds of fruit: (A), (B) and (C) are 
radial, tangential and diagrammatic cross-sectional views of a ray 
fruit; (D) and (E) are radial and diagrammatic cross-sectional 
views of a central disc fruit. Peripheral disc fruit are larger and 
angular ovate in cross-section. 

12 Rhodora [Vol. 76 

curate in light of the findings of my investigations. The 
previously unreported heterocarpic fruit condition is also 
described (Figure 5). A complete list of specimens exam- 
ined in the study is given in Semple (1972b) and only rep- 
resentative specimens are cited here. 

Xanthisma DC., Prodr. 5:94-95. 1836. 
Centauridium T. & G., Fl. N. Amer. 2:246. 1842. 

Taprooted annuals, rarely biennial and flowering the 
second season; branches ascending, few to many. Leaves 
alternate, the upper sessile, the lower terminally serrate; 
basal leaves petioled, lobed or pinnatifid to bipinnatifid. 
Heads solitary; involuere turbinate to hemispheric or cam- 
panulate; involucral bracts broad in 3-4 series, imbricate, 
stiff, the margins pale; ray florets pistillate, fertile, yellow; 
dise florets perfect, fertile, yellow. Fruit heterocarpic with 
pappus bristly and margins finely serrate; both fruit kinds 
with fine ascending white hairs, straw-colored when mature 
and darkening with age. Receptacle slightly convex, with 
subulate chaff reticulately dispersed between the fruits. 

The genus is monotypic and is endemic to the central 
southwestern areas of the United States. 

Xanthisma texanum DC., Prod, 5:95. 1836. 
Lectotype: Texas; Bexar County, in woods near Medina, 
May 1828, Berlandier 2039d (G!). Isolectotypes; G! (4), 
GH! (1), NY! (1), and us! (2). 

Herbs 2-7 (rarely more) dm tall, branching near the 
middle to ground level from a main stem 0.1-3 dm long, or 
rarely acaulescent. Upper leaves 1-2 cm long, linear, entire; 
basal leaves 5-8 cm long, ovate to obovate in outline, inter- 
grading above. Heads solitary, not crowded; involucre 
5-10 mm high, involucral bracts 1.5-4 mm broad, rarely 
recurved, margins pale (the lateral margins very rarely 
red), the central portion green, reticulate venation pro- 
nounced, the inner-most bracts chartaceous, the outer-most 
linear grading into the peduncle bracts; ray florets 10-30, 
the straps 8-15 mm long, yellow adaxially, pale yellow 

1974] Xanthisma — Semple 13 

abaxially (rarely red-tinted abaxially) ; disc florets many 
40-150, slightly ampliate to tapering, the lobes short. Ray 
floret cypselas curved obpyramidal, triangulate in cross- 
section, the pappus grading from short linear to long 
basally broad scales; disc floret cypselas obovate radially 
and narrowly-so tangentially, the central fruits rhombic 
in ercss-section, the outer obtrullate, the pappus in two 
whorls, the inner members ca. 6 mm long, linear, 8-15, the 
outer members 3 mm long or less many. 

The species is characterized by the following combination 
of characters: heterocarpic fruit with a pappus of bristly 
seales only, fruit pubescence of long white ascending hairs, 
receptacle slightly convex with a persistent reticulate net- 
work of subulate scales. 

The lectotype of the species was chosen from five speci- 
mens seen on microfilm only (1DC micro edition of Candolle 
Prodromi Herbarium). Since all five specimens are similar 
in quality, the specimen with the complete label giving col- 
lection number, date and location, and floral color (fourth 
from the left on the microfilm) is designated lectotype. 
Torrey (1859) first noted the typographic error in the Pro- 
dromus, which cited the type collection as Berlandier 2639 
rather than Berlandier 2039. 

Key to Infraspecific Taxa 

I. Involucral bracts shorter than wide above the widest 
part, 1.5-3.5 mm wide and 1-2 mm long above the widest 
We eS, ur. spe Feats d yr g e raa subsp. texanum (1) 
II. Involucral bracts rounded with little or no flare 

1 mm from the apex, occasionally recurved. 
i gu ck na ease esse es ewer’ var. texanum (2) 
II. Involueral bracts obtusely pointed, the deltoid por- 
tion 2-3 mm wide and 1.5-2 mm long above the 
widest part. ............-...-- var. orientale (3) 
III. Lateral margins of bracts whitish. .......... 
JI NRORURUIOPNBA TO ROS .... f. orientale: (38) 
III. Lateral margins of bracts red. f. rubrum (3b) 

14 Rhodora [Vol. 76 

I Involucral bracts longer than wide above the widest 
part; deltoid portion 2.5-5 mm wide and 3.5-8 mm long 
in bracts of the middle series. .. ssp. drummondii (4) 

1. Xanthisma texanum spp. texanum 

Xanthisma texanum var. berlandieri Gray, Smiths. 
Contrib. Knowl. 3(5) : 98. 1852. 

Xanthisma berlandieri (Gray) Small, Fl. SE. U.S. p. 
1184. 1903. 

This subspecies includes the type collection of X. texanum 
and is distinguished by small involucral bracts with rounded 
or obtusely pointed apices. Two varieties are recognized 
within the subspecies. 

2. Xanthisma texanum ssp, texanum var. texanum 

The type variety of the species is distinguished by its 
small apically rounded involucral bracts 1.5-2 mm wide, 
appressed or rarely recurved; generally 30-35 bracts are 
visible without removing the outer series. The variety is 
endemic to the Nueces Plains of southern Texas from San 
Antonio to the south and west. East of San Antonio this 
variety forms hybrids with subspecies drummondit. 

Charlotte, 4 May 1919, Schulz 39 (Us). Bexar Co., San Antonio, 
4 June 1948, Burr 21 (NY, TEX), in woods near the Medina River, 
July 1829, Berlandier 629 (GH, MO, NY, US). Dimmit Co., Big Wells, 
21 April 1945, Shinners 7,400 (NY, OKLA, SMU, TEX), in front of 
cemetery, 6 May 1964, Turner 4,996 (TEX). Frio Co., Dilley, 6 mi. 
W, 8 July 1970 Lewis 7,634 (Mo). La Salle Co., Artesia Wells, 3 mi. 
W., 7 April 1963, Dickey 170 (SMU, TEX). Medina Co., Farm Rd. 
1343, 4.4 mi S of U.S. Hwy. 90, 29 June 1972, Semple & Shea 720 
(MO) ; Devine, 3 mi. SW, 1 May 1954, Tharp & Turner 3,439 (OKLA, 
TEX). Zavala Co., Batesville, 6 mi. E, 6 May 1964, Turner 5,002 

9. Xanthisma texanum ssp. texanum var. orientale Semple, 
var. nov. 
Involucri bracteae parvae manifeste depresse ovatae, 
apice obtuse, cuspidatae, interdum marginibus lateralibus 

1974] Xanthisma — Semple 15 

HOLOTYPE: Texas: Hidalgo Co.: 13.5 mi. N. of Edinburg on 
U. S. Hwy. 183, 8 July 1970, W. Lewis 1642 (Mo). Iso- 
types: four, to be distributed. 

The involucral bracts are small and pronouncedly shield- 
shaped, occasionally with red lateral margins. The apices 
are obtusely cuspidate. 

This variety is distinguished by the 2-3.5 mm wide in- 
volucral bracts having obtuse, cuspidate apices. The short 
stem habit is also frequently encountered. Hybridization 
with var. texanum is common, but is unknown with ssp. 
drummondii. The name is taken from its distribution in 
the eastern half of the range of ssp. texanum. Two forms 
are recognized. 

3a. Xanthisma texanum var. orientale Semple f. orientale 
The type form is the most common form of the variety. 

Aransas Refuge, 29 Sept. 1944, Cory 45, 895 (Ny, TEX). Bee Co., 

Beeville, 21 June 1935, Drushel 9,941 (NY). Brooks Co., Hebbron- 

ville, 15 mi. E, 25 Nov. 1962, Dohnke T (sMU). Calhoun Co., Port 

Lavaca, 14 June 1953, Johnston 53.280.157 (OKLA, TEX). Goliad Co., 

Goliad, 15.9 mi. S, 9 July 1970, Lewis 7,655 (Mo). Hidalgo Co., Rio 

Grande Valley, 8 Aug. 1942, Walker 70 (KSU, TEX). Jim Hogg Co., 

Hebbronville, 8 mi. E, 8 July 1970, Lewis 7,638 (MO). Jim Wells Co., 

Premont, 12 mi. N, 24 Nov. 1954, Johnston 542,144 (TEX). Kenedy 

Co., King Ranch, Norias Div. 24 Nov. 1953 Johnston 53.280.154 

(OKLA, TEX.) Kleberg Co., Kingville, 1940, Sinclair s.n. (TEX). 

Refugio Co., 5 mi. S of county line on U. S. Hwy. 77A, 12 June 

1971, Semple 581 (Mo). San Patricio Co., Aransas Pass, 24 May 

1922, Schulz 868 (TEX, US). Near San Patricio, May 1834, Berlandier 

2578 (GH, MO, NY). Webb Co., Mirando City, 31 Aug. 1936, Baird 

s.n. (NY). Willacy Co. Yturria Station, 6 Aug. 1924, Runyon 664 

(TEX, US). 

3b. Xanthisma texanum var. orientale f. rubrum Semple, 
forma nov. Involucri bracteae cum marginibus later- 
alibus rubris. 

HOLOTYPE: Texas: San Patricio Co., 0.5 mi. E. of Ingleside 
on Texas Hwy. 361, 13 June 1971, Semple 602 (Mo). 
Isotype: one to be distributed. 

This form is distinguished from the type form by having 
bracts with red lateral margins. 

16 Rhodora [Vol. 76 

Aransas Refuge, 29 Sept. 1944, Cory 45,895 (Mo). Goliad Co., 
General Zaragoza State Park, 1 mi. S, 12 June 1971, Semple 579 
(mo). Kenedy Co., Raymondville, 16 mi. N, 2 Dec. 1945, Cory 
51,493 (us). Willacy Co., Port Mansfield, 0.5 mi. W of beach by 
garbage dump, 9 July 1970, Lewis 7,644 (MO). 

4. Xanthisma texanum DC. ssp. drummondii (T. & G.) 
Semple, stat. nov. 
Centauridium drummondii T. & G., Fl. N. Amer. 2: 462. 
SYNTYPES: Texas, north of San Antonio, Riddell s.n. (NY) ; 
Texas, exact locality unknown, Nov. 1835, Drummond 
227 (GH!, NY!). 
LECTOTYPE: Drummond 227 (GH). Isolectotypes: GH, NY. 

Xanthisma texanum var. drummondii (T. & G.) Gray, 
Smiths. Contrib. Knowl. 3(5) : 98. 1852. 

Xanthisma drummondii (T. & G.) Hooker f., Curtis’ 
Bot. Mag. 33: t. 6275. 1877. 

Subspecies drummondii is distinguished by its involucral 
bracts, which are wider than those of ssp. texanwm and are 
distinctly lanceolate. Populations occur in much of Texas 
north of San Antonio, and in Oklahoma and eastern New 

Drummond 227 (GH) was selected lectotype because it is 
the largest and most complete specimen. The Riddell col- 
lection cited by Torrey & Gray (1842) is fragmentary and 
a putative hybrid between the two subspecies. 

velt Co., Elida, 5 mi. NE, 28 Sept. 1965, Anderson 3,004 (KSU). Lingo, 
1.2 mi. N, Semple & Shea 705 (MO). OKLAHOMA: Beckham Co., 
Carter, 5 mi. S, 5 July 1965, Thomas 79 (OKLA). Caddo Co., Cement, 
27 June 1936, Demaree 18, 133 (NY). Comanche Co., Cache, June 
1927, Ortenburger s.n. (U.S.) Creek Co., Drumright, 10 June 1935, 
Fry 8 (OKLA). Custer Co., Clinton, 28 June 1966, Ganz s.n. (ARIZ. 
st. U.). Grady Co., Verdun, 25 June 1963, Pearce 885 (OKLA, SMU). 
Greer Co., Granite, 17 June 1939, Grawmanns 85 (OKLA). Harmon 
Co., Vinson, Shinners 31,644 (sMU). Jackson Co., Duke, 1.7 mi W. 
15 July 1970, Flyr s.n. (MO). Kay Co.; 26 July 1898, White s.n. 
(us). Kiowa Co. Mt. Park, 23 June 1913, Stevens 1,293 (OKLA). 

1974] Xanthisma — Semple 17 

Lincoln, 23 Aug. 1895, Blankenship 17,334 (wis). Logan Co., Guthrie, 
12 July 1916, Keyser 6,036 (Ny). Osage Co., 8 mi. S of Kans. state 
line on Okla. Hwy. 99, 24 June 1972, Semple & Shea 667 (MO). 
Oklahoma Co., Oklahoma City, 27 May 1937, Waterfall 582 (OKLA, 
NY). Pawnee Co., Pawnee, 8 mi. E, 8 June 1934, Brodell s.n. (TEX). 
Payne Co., Mulhall, 1.1 mi. N, 25 June 1972, Semple & Shea 674 
(MO). Stephens Co., Duncan, 8 mi. N, 27 May 1960, Waterfall 15,911 
(OKLA). Washita Co., 2.2 mi. E of county line on Okla. Hwy. 152, 
20 Aug. 1970, Semple & Love 264 (Mo). TEXAS: Andrews Co., 
Andrews, 4 mi. W, 28 May 1956, Turner 3,957 (TEX). Archer Co., 
Mankins, 2 mi. NE, 8 June 1956, Shinners 23,758 (SMU). Baylor Co., 
Seymour, 4 Aug. 1955, Shinners 20,776 (SMU). Blanco Co., Silverton, 
Farm Rd. 145, 26 June 1972, Semple & Shea 699 (M0). Burleson Co., 
Caldwell, 14 June 1971, Semple 625 (Mo). Burnet, 8 mi. W, 19 May 
1955 Turner & Johnston 2,480 (TEX). Callahan Co., Baird, Aug. 1882, 
Letterman 26 (Mo, US). Childress Co., Childress, Biology Class C. 
High School 12 (TEX). Coke Co., Robert Lee, 3 mi. S, 4 Oct. 1968. 
Gary 192 (TrC). Comanche Co. Comyn, Theney School s.n. (TEX), 
Dallas Co., Dallas, near Comanche Peak, Aug. 1877, Reverchon s.n. 
(vs). Dewitt Co., western part of Co., 20 July 1941, Reidel s.n. 
(TEX). Erath Co., 1922, Gaugh 29 (vs). Fisher Co., Rotan, 10 Sept. 
1933, Brooker s.n. (TEX). Gaines Co., Lamesa, 15.1 mi. W, 14 Sept. 
1946, Whitehouse 16,788 (NY, SMU). Gillespie Co., Cherry Spring, 
18??, Jermy 639 (Mo). Gonzales Co., Cost, 11.5 mi. SW, 29 June 
1972, Semple & Shea 730 (Mo). Hall Co., Turkey, 5 mi. S, 12 June 
1950, Tilton T.550.604 (TTC). Hood Co., 6 Aug. 1877, Reverchon 
1368 (KsU. NY, US). Howard Co. Big Spring, 22 mi. S, 27 April 
1963. Dallas 64 (OKLA). Kerr Co.. 6 June 1929, Whitehouse 7.056 
(TEX). Limestone Co.. Kosse, 3.5 mi. N, 14 June 1971, Semple 626 
(MO). Lubbock Co., Lubbock, Demaree 7,715 (MO, TEX, TTC. US). 
McCullogh Co., 1 mi. S of Colorado R., 28 June 1972, Semple & Shea 
716 (MO). Navarro Co., Corsicana, 21 June 1872, Hall 311 (GH, MO, 
NY, Us). Robertson Co., Calvert, 8 May 1904, Tyler s.n. (US). Tar- 
rant Co. Lake Fort Worth, 16 July 1923, Ruth 742 (GH, KSU, US, 
wis). Travis Co. Austin, 9 June 1908, Biltmore s.n. (US). Van 
Zandt Co., Edom, 3 July 1939, Ball 7 (sMv). Washington Co., 20 
May 1938, Brackett s.n. (TEX). Young Co. Belknap, 6 May 1858, 
Hayes 397 (NY). 


Braun, E. L. 1950. Deciduous Forests of Eastern North America. 
Blakiston Co. Philadelphia. 

Bray, W. L. 1906. Distribution and Adaptation of the Vegetation 
of Texas. Bull. of U. of Tex. No. 82, Sci. Ser. No. 10. 

18 Rhodora [Vol. 76 

CANDOLLE, A. P. DE. 1836. Prodromus 5: 94-95. 

CLAUSEN, R. T. 1941. On the terms “subspecies” and “variety.” 
Rhodora 43: 157-167. 

CORRELL, D. S. & M. C. JoHNSTON. 1970. Manual of the Vascular 
Plants of Texas. Texas Research Foundation Press. Austin. 

Du RErTZ, G. E. 1930. The fundamental units of biological tax- 
onomy. Svensk. Bot. Tidskr. 24: 333-428. 

EHRENDORFER, F. 1968. Geographical and ecological aspects of 
infraspecific differentiation. In V. H. Heywood, ed. Modern 
Methods of Plant Taxonomy. Academic Press. New York. 

FENNEMAN, N. M. 1931. Physiography of the Western United 
States. MeGraw-Hill Book Co. New York. 

FERNALD, M. L. 1940. Some spermatophytes of eastern North 
America. Rhodora 42: 239-278. 

FoscuE, E. J. 1932. The natural vegetation of the lower Rio 
Grande Valley, Texas. Field & Lab. 1: 25-30. 

GOULD, F. W. 1962. Texas Plants— a checklist and ecological 
summary. Texas A. & M. — Tex. Agr. Ex. Sta. College Station, 

GRAY, A. 1852. Plantae Wrightiana  Texano — Neo-Mexicanae. 
Part I. Smiths. Contrib. Knowl. 3: 98. 

Hooker, J. D. 1877. Curtis’ Bot. Mag. 33: t.6275. 

OETKING, P. F. 1959. Geological Highway Map of Texas. Dallas 
Geological Soc. Dallas. 

Rarsz, E. 1957. Land forms of the United States. Jn Oetking, 
P.F. 1959. Geological Highway Map of Texas. Dallas Geological 
Soc. Dallas. 

SELLARD, E. H., W. S. ADKINS & R. B. PLUMMER. 1966. Geology of 
Texas. Vol. I. Stratigraphy. U. of Tex. Bull. No. 3232. 

SEMPLE, J. C. 1972a. Behavior of B-chromosomes in Xanthisma 
texanum DC.: a non-random phenomenon. Science 175: 666. 

1972b. Cytology, Flavonoid Chemistry and System- 
aties of the Texas Sleepy Daisy Xanthisma texanum DC. (As- 
teraceae). Ph.D. Dissertation. Washington University. St. Louis. 

SMALL, J. K. 1903. Flora of the Southeastern United States of 
America. Pub. by Small. New York. p. 1184. 

SMITH, E. B. & H. M. Parker. 1971. A biosystematic study of 
Coreopsis tinctoria and C. cardaminefolia. Brittonia 23: 161-170. 

STEENIS, C. G. G. J. VAN. 1957. Specific and infraspecific delimina- 
tion. Flora Malesiana Ser. 1, 5: clxvii-eexxxix. 

THARP, B. C. 1926. Structure of the Texas Vegetation East of the 
98th Meridian. U. of Tex. Bull. No. 2606. 

Torrey, J. 1859. In Emory, W. H., ed. Report on the United States 
and Mexican Boundary Survey. Nicholson Press. Washington. 
Vol. II. pp. 80-81. 

1974] Xanthisma — Semple 19 

. & A. GRAY. 1842. A Flora of North America. Vol. II. 
facsimile ed., 1969. Hafner Pub. Co. New York. p. 246. 
VALENTINE, D. H. & A. LóvE. 1958. Taxonomic and biosystematic 
categories. Brittonia 10: 153-166. 
VISHER, S. S. 1954. Climatic Atlas of the United States. Harvard 

U. Press. Cambridge. 

ST. LOUIS, MO. 63110 

flowered form of Utricularia purpurea Walter was found 
while conducting research in a small pond in New Hamp- 
shire. This form was extremely abundant forming exten- 
sive mats on the surface of this shallow pond. None of 
the normal purple-flowered plants were observed. Plants 
commonly associated with the white form were Utricularia 
vulgaris L., Utricularia intermedia Hayne and Najas flexilis 
Rostk. & Schmidt. The pond had a pH of 6.5 and methyl 
orange alkalinity reading of 8.0 mg/1. 

The following form is described. Utricularia purpurea 
Walter forma alba Hellquist, forma nova. differt forma 
purpurea quantum flores albos non purpureos habet. TYPE: 
NEW HAMPSHIRE: CARROLL COUNTY: small unnamed pond 
southeast of Dorr Pond on N.H. Route 153 at the settlement 
of Woodman, Town of Wakefield (U.S.G.S. quadrangle 
Newfieid, Me.-N.H.), C. B. Hellquist 8935 (NHA). 

The holotype has been placed at the University of New 
Hampshire Herbarium (NHA). Isotypes are in the her- 
barium of the New England Botanical Club (NEBC) and 
Boston State College Herbarium. 





The Longleaf Pine of the southeastern United States has 
been spared the numerous changes in scientific name that 
have cast confusion over many of our other tree species. 
From pioneer days the dominant practice, both among 
botanists and foresters, has been to apply the name Pinus 
palustris Mill. to the Longleaf. Prior to the 1940's the 
only author of significance to do otherwise was J. K. Small 
(1933) who interpreted P. palustris Mill. to be the northern 
variety of slash pine, P. elliottii Engelm. Small’s Manual 
of the Southeastern Flora, however, achieved only regional 
impact, and his use of P. australis Michx. f. for the Long- 
leaf did not gain a wide following. 

A much stronger impetus for associating the Longleaf 
with Pinus australis was generated in 1948 by M. L. Fer- 
nald who advocated this name in two detailed and strongly 
worded articles (Fernald & Schubert, 1948; Fernald, 
1948). Fernald's position was reinforced by the publica- 
tion of his monumental reworking of Gray's Manual of 
Botany (1950), and was further entrenched when the inde- 
pendent H. A. Gleason adopted the same name in his New 
Britton and Brown Illustrated Flora (1952) as did A. 
Cronquist in their companion field guide, the Manual of 
Vascular Plants of Northeastern United States and Adja- 
cent Canada (Gleason & Cronquist, 1963). These three 
publications totally dominate the field of regional floras in 
the Northeast, and their consistent use of Pinus australis 
inevitably suggests that this name is supported by general 
professional agreement and by nomenclatural legitimacy. 

Yet in publications other than these or their derivatives, 
the Longleaf remains known by the name Pinus palustris 

"This paper is Florida Agricultural Experiment Station Journal 
Series No. 4775. 


1974] Longleaf Pine — Ward 21 

Mill. This is the name found in state or district floras 
such as the Manual of the Vascular Flora. of the Carolinas 
(Radford et al., 1968), the Manual of the Vascular Plants 
of Texas (Correll & Johnston, 1970), and A Flora of 
Tropical Florida (Long & Lakela, 1971), as well as in the 
more basic modern treatments of the genus by Mirov 
(1967) and by Little & Critchfield (1969). The U. S. Forest 
Service, following the guidance of its dendrologist, E. L. 
Little, has remained steadfast with P. palustris, and the 
state forest services have done likewise. 

The rationale behind Fernald's heterodox displacement 

of Pinus palustris was his view that Philip Miller (1768), 
superintendent of the Chelsea Physic Garden and British 
author of a series of widely used and authoritative horti- 
cultural dictionaries, did not have the Longleaf when he 
described an American tree under this name. The original 
description contained phrases that do not apply to the 
stately and versatile Longleaf: “I have been informed that 
they grow to a height of twenty-five or thirty feet. 
I have not heard the wood was of any use but for fuel." 
However, Miller reported these statements as hearsay, not 
as observed fact, while his description also contained such 
passages as ‘“‘Pine-tree with the longest leaves growing by 
threes out of each sheath" and “Their leaves are a foot or 
more in length, growing in tufts at the end of the branches, 
so have a singular appearance." These phrases were given 
without qualification and form a brief but unmistakable 
description of the Longleaf, 

Fernald further relied on the habitat of Pinus palustris 
reported by Miller, “swamps in many parts of North 
America," as excluding the Longleaf. He proposed instead 
that Miller may have had the Loblolly Pine, P. taeda L. 
Actually however, although the Longleaf usually occurs on 
dry well-drained soils, it not uncommonly is found associated 
with the Slash Pine in the extensive level undrained swamps 
known as flatwoods and throughout its range probably 
occurs as frequently in wet soils as Fernald's proposed 
substitute, the Loblolly. Of couse, Miller's use of an epithet 

22 Rhodora [Vol. 76 

meaning “swampy”, even if it were totally unsuitable, 
would not be justification for discarding his P. palustris; 
Article 62 of the International Code of Botanical Nomen- 
clature (Stafleu, 1972) provides that: “a legitimate name 
or epithet must not be rejected merely because it is in- 
appropriate . . ." 

No information is available from specimens preserved 
by Miller; Dr. Schubert examined the Miller collections of 
the British Museum and was unable to find a specimen that 
could be considered a type. Fernald, by correspondence, 
did locate a specimen he presumed to be of P. taeda that 
Miller may have seen and that someone, possibly Miller, 
had noted as “palustris.” It seems improbable that Miller 
would have so misidentified P. taeda since this species was 
well known to him and was treated in some detail in the 
several editions of his “Dictionary.” 

Fernald did demonstrate at some length that the name 
Pinus palustris was not clearly understood and was even 
misused by Miller's contemporaries and other early workers, 
but this again does not provide justification for discarding 
the name. There seems to be essentially no sound rationale 
for interpreting this name to mean anything other than, 
in the conventional usage, the Longleaf Pine. 

In the event, moreover, that Miller's brief but clear 
diagnosis should be considered ambiguous, no alternative 
name is available. Pinus lutea Walter, whimsically sug- 
gested by Fernald, is probably P. taeda. Pinus australis 
Michx. f. is unmistakably the Longleaf, and was selected 
as the correct name for this species by Small and by 
Fernald. Little (1948), however, accurately pointed out 
prior to the publication of Fernald's first paper on the 
subject, that Article 63 (then Article 60) of the Inter- 
national Code makes this name illegitimate since it was 
superfluous when published. Not only did Michaux (1810) 
cite P. palustris as a synonym, but he specifically explained 
that he was replacing this name with P. australis, using 
an epithet (“southern”) that he thought more appropriate. 
It is precisely such actions as this that Article 63 is de- 

1974] Longleaf Pine — Ward 23 

signed to prevent, and Fernald's protestations that Michaux 
was “definitely defining a new species" are contradicted by 
Michaux' own words, in translation: *I have thought like- 
wise that the specific name ‘australis’ was preferable to 
that of ‘palustris’, under which this species has been de- 
scribed by botanists; for this last gives an absolutely false 
idea of the nature of the soil where this tree grows.” 

The only other specific name apparently ever given the 
Longleaf was Pinus longifolia Salisb., but here again the 
epithet used was superfluous, for Salisbury (1796) was un- 
abashedly providing a substitute name for P. palustris and 
thereby formed an illegitimate and unusable combination. 

The person who would apply a scientific name to the 
Longleaf Pine, therefore, has the choice, should he wish to 
follow the Code, of either accepting Miller’s brief descrip- 
tion of Pinus palustris as adequate, or of discarding it as 
confused and coining and publishing a name of his own 
creation. Neither P. australis Michx. f. nor P. longifolia 
Salisb. may be legitimately used for this tree, and prudence 
and practicality, as well as historical precedent, indicate 
strongly the advisability of retaining the Longleaf Pine 
under the name Pinus palustris Mill. 


CORRELL, D. S. & M. C. JOHNSTON. 1970. Manual of the Vascular 
Plants of Texas. Texas Research Found. 1881 p. 

FERNALD, M. L. 1948. The confused bases of the name Pinus 
palustris. Rhodora 50: 241-249. 

. 1950. Gray's Manual of Botany, 8th ed. Ameri- 

can Book Co. 1632 p. 

, & B. G. SCHUBERT. 1948. Studies of American 
types in British herbaria. Rhodora 50: 181-190. 

GLEASON, H. A. 1952. The New Britton and Brown Illustrated 
Flora. Lancaster Press. Vol. 1. 

,& A. CRONQUIST. 1963. Manual of Vascular Plants 
of Northeastern United States and Adjacent Canada. Van Nos- 
trand Co. 810 p. 

LITTLE, E. L. 1948. Notes on nomenclature of trees. Phytologia 
2: 457-458. 

24 Rhodora [Vol. 76 

, & W. B. CRITCHFIELD. 1969. Subdivisions of the 
genus Pinus (pines). U. S. Dept. of Agric. For. Ser. misc. publ. 
1144. 51 p. 

Lonc, R. W., & O. LAKELA. 1971. A Flora of Tropical Florida. 
Univ. of Miami Press. 962 p. 

MicHAUX, F. A. 1810. Histoire des Arbres Forestiers de l'Amerique 
Septentrionale. 1: 64. 

MiLLER, P. 1768. The Gardeners Dictionary, 8th ed. London. 

Mirov, N. T. 1967. The Genus Pinus. Ronald Press. 602 p. 

RADFORD, A. E., H. E. AHLES, & C. R. BELL. 1968. Manual of the 
Vascular Flora of the Carolinas. Univ. N. Car. Press. 1183 p. 

SALISBURY, R. A. 1796. Prodromus Stirpium in Horto ad Chapel 
Allerton Vigentium. London. 

SMALL, J. K. 1933. Manual of the Southeastern Flora. The 
author, New York. 1554 p. 

STAFLEU, F. A. 1972. International Code of Botanical Nomencla- 
ture. Utrecht. 426 p. 


1972, Mr. Richard Fackovec, a student in an aquatic 
botany course at Southampton College, brought to our 
attention a specimen of Taxodium distichum that he col- 
lected along the shore of Long Pond in Sag Harbor, New 
York. We returned to investigate the region in November 
and found eight bald cypress seedlings, ranging in height 
from one to 214 meters and growing close to shore in 
0.5-0.7 meters of water. The four smallest trees (approxi- 
mately one meter tall) are growing close together at the 
southern end of the pond at the edge of a large marsh. The 
other cypress trees are located at the southern end of the 
pond also, but along the western shore. The two largest 
trees (215 and 114 meters tall) are at the edge of a large 
stand of Typha latifolia. Most of the remaining shoreline 
of Long Pond consists of very dense vegetation composed 
primarily of shrubs and small trees (e.g. Myrica gale, 
Decodon verticillata, Cassandra calyculata, Vaccinium 
corymbosum, Nyssa sylvatica, Acer rubrum). It is within 
this shrub zone that we found two additional bald cypress 
seedlings. One had become encompassed so completely by 
encroaching shoreline vegetation that we removed it; sub- 
sequently, it has been replanted along the shore of a pond 
in the Morton Wildlife Refuge, a Federal Wildlife Preserve 
in Noyac, New York. 

We have communicated with Dr. Richard Stalter, plant 
taxonomist at St. Johns University in Jamaica, New York, 
regarding the northern distribution of Taxodium distichum. 
As far as we have been able to determine, this is the first 
record of a natural population of bald cypress in New 
York State. The closest naturally-occurring populations 
of Taxodium are in Delaware, where there are a few large 
colonies (Stalter, personal communication). There was, 
at one time, a colony in New Jersey, but it has been un- 
known in that state for several years (Stalter, personal 


26 Rhodora [Vol. 76 

Accompanied by Dr. Stalter, we returned to Long Pond 
in December in an unsuccessful effort to locate mature 
bald cypress trees. We share the opinion with Dr. Statler 
that this colony of Taxodium distichum represents a natu- 
ralized population. The presence of a few bald cypress 
seedlings in relatively inaccessible portions of the shore- 
line strongly suggests that they were not planted. The 
apparent absence of large bald cypress in the area leaves 
unanswered the question as to how the seedlings became 
established; hopefully, further investigation will afford us 
an answer. 

The voucher specimen collected by Mr. Fackovec is de- 
posited in the Southampton College Herbarium. 






Grand Beach Provincial Park lies approximately 57 miles 
north of Winnipeg. The entire park, which covers 59,000 
acres, is in the Manitoba Lowlands Section of the Boreal 
Forest Region (Rowe, 1959). Three miles of the area 
fronts on Lake Winnipeg. 'The underlying bedrock is 
palaeozoic limestone, and the overlying beds are modified 
till, or lacustrine clays deposited in glacial Lake Agassiz. 
Most soils have a high lime content. 

On the flat, poorly drained areas, black spruce (Picea 
mariana (Mill. BSP.) and tamarack (Larix laricina (Du 
Roi) K. Kich) forest prevail, while on the better drained 
alluvial strips bordering the rivers and creeks, white spruce 
(Picea glauca (Moench) Voss), aspen (Populus tremuloides 
Michx.) and balsam poplar (Populus balsamifera L.) pre- 
dominate. On the low, well-drained ridges, jack pine (Pinus 
banksiana Lamb.) is dominant, with aspen, paper birch 
(Betula papyrifera Marsh.) and some bur oak (Quercus 
macrocarpa Michx.) present. 

No bryophytes have previously been reported for the 
park. In the present study, several sites were visited, 
mostly in the vicinity of 96°35’W and 50°33’N. The sites 
were chosen as representative of the major vegetation types 
in the area, and the following seven, visited in July, 1972, 
are reported as representative of the area studied. Several 
of the vascular collections could be determined to genus 
only, because of lack of flowering or fruiting material. 

1. Sand dunes and dune slacks behind the East Beach. 
Dominant shrubs were Salix spp., especially in the slacks, 
with Prunus pumila L. conspicuous on the drier areas. 
Other prominent vascular plants on the dry areas included 
Carex spp. Rhus radicans L. var. rydbergii (Small) 


28 Rhodora [Vol. 76 

Rehder, and Artemisia spp. Total plant cover was rather 
thin (about 20%) except in the slacks, and most bryophyte 
cover was on wet litter in the slacks. Small turf mosses 
such as Bryum spp. were, however, common in dry habi- 

2. Aspen-dominated forest just s. of the parking lot at the 
East Beach. Most mosses were from tree bases. 

3. Mixed mature aspen — jack pine forest on w. side of 
the road n. of main park entrance. 

4. Black spruce bog with some tamarack and white cedar 
(Thuja occidentalis L.). The understory was dominated by 
Ledum groenlandicum Oeder and the terrain consisted of 
pools and Carex hummocks. Just s. of Highway 12, 14 mile 
from its junction with Highway 59. 

5. Open jack pine with some birch, on s. side of Highway 
12, 2 miles from its junction with Highway 59. 

6. Tamarack — black spruce bog on e. side of Highway 59, 
2 miles n. of its junction with Highway 12. Drier than 
site 4, and with a tall shrub layer of Alnus sp., as well as a 
short shrub layer dominated by Ledum groenlandicum. 
Deadfall was abundant. 

7. Mature, open jack pine forest on w. side of Highway 59 
4 miles n. of its junction with Highway 12. There was an 
open short shrub and herb layer dominated by Vaccinium 
angustifolium Ait., Chimaphila umbellata (L.) Bart. and 
Amelanchier alnifolia Nutt., with Linnaea borealis L. and 
Vicia americana Muhl. prominent. The forest floor was 
dominated by Plewrozium schreberi and Dicranum spp., 
especially D. polysetum. The many large humus-covered 
boulders present all had an extensive, species-rich bryo- 
phyte cover. Numbers in the species list refer to the sites 
described above. 

Taxonomic and field work were shared equally by both 
authors. Voucher specimens have been deposited in the 
authors’ own herbarium and at the University of Winnipeg. 
Nomenclature for Sphagnobrya and Eubrya follows Crum, 
Steere, and Anderson (1965) with modification according 
to Crum (1971). Hepatic nomenclature follows Schuster 

1974] Bryophytes — Stringer & Stringer 29 

(1969), supplemented by Schuster (1953) with abbrevia- 
tions of authorities amended to conform with the list of 
Sayre, Bonner and Culberson (1964). Vascular plant no- 
menclature follows Scoggan (1957). 


Calypogeia muelleriana (Schiffn.) K. Miill. (6). Three 
collections, two on humus mixed with well-rotted wood, 
and one on a rotten stump with Lophocolea heterophylla, 
Cephalozia media and Hypnum pallescens. Only once pre- 
viously recorded for the province (Longton, 1972). 

Cephalozia media Lindb. (6). One collection, on a rotten 
stump. | 

Cephaloziella rubella (Nees) Douin (4). On humus over 
rotten wood, with Amblystegium juratzkanum, Chiloscy- 
phus pallescens and Thuidium recognitum. 

Chiloscyphus pallescens (Ehrh.) Dum. (4). Only once pre- 
viously recorded for the province, in Spruce Woods Pro- 
vincial Park (Stringer & Stringer, a, in preparation). 

Chiloscyphus polyanthus (L.) Corda (3, 4). On bark at the 
base of an aspen tree (3) with Hypnum pallescens, and 
on humus with Distichium inclinatum. Recorded previ- 
ously only in Spruce Woods Provincial Park (Stringer & 
Stringer, in preparation). 

Lophocolea heterophylla (Schrad.) Dum. (3, 6, 7,). On 
humus or rotten wood, often with Pohlia nutans or Hyp- 
num pallescens. Previously recorded only in the Spruce 
Woods area by Bird (1969) and by Stringer & Stringer 
(in preparation). 

Lophocolea minor Nees (6). On thick humus (dead Aula- 
comnium palustre), with Pohlia nutans and Eurhynchium 
pulchellum. Not previously recorded for the province. 

Ptilidium ciliare (L.) Nees (5, 6, 7,). On loose humus or 
litter with Dicranum spp. and Pleurozium schreberi. One 
collection (7) on loose conifer needles over rock, with 
Hedwigia ciliata. 

Ptilidium pulcherrimum (Web.) Hampe (7). One collec- 
tion, on fallen birch bark. 

30 Rhodora [Vol. 76 

Radula complanata (L.) Dum. (7) One collection, on packed 
humus at the base of a jack pine tree. 

Riccardia pinguis (L.) S. Gray (4). On thick, moist humus 
mixed with silt, with Myurella julacea, Amblystegium 
juratzkanum and Platydictya jungermannoides. Previ- 
ously recorded only in the Spruce Woods area by Bird 
(1969) and Stringer & Stringer (in preparation). 

Sphagnum capillaceum (Weiss) Schrank (6). In deep hum- 
mocks, with Pleurozium schreberi. 


Amblystegium juratzkanum Schimp. (1, 2, 4, 5, 6, 7). On 
rotten wood or humus, especially at tree bases, where it 
was often found with Pylaisiella polyantha and Brachy- 
thecium salebrosum. 

Amblystegium varium (Hedw.) Lindb. (1, 2, 6). On thick 
humus and rotten wood, with Brachythecium spp. and 
Mnium cuspidatum. 

Aulacomnium palustre (Hedw.) Schwaegr. (4, 5, 6). On 
thick humus, in deep pure sods, or with Pleurozium 
schreberi. One collection (5) on rotting birch bark. 

Brachythecium campestre (C. Müll.) B. S. G. (5, 6, 7). On 
humus, usually over well-drained sandy soil. Previously 
recorded in Manitoba only in the Winnipeg and Spruce 
Woods areas by Stringer & Stringer (1973) and Stringer 
& Stringer (in preparation). 

Brachythecium rutabulum (Hedw.) B. S. G. (1, 2, 5, 6, 7). 
On humus or rotten wood in moist habitats. Previously 
recorded only by Mueller-Dombois (1964), Longton 
(1972), and by Stringer & Stringer (in preparation), in 
the Spruce Woods area. 

Brachythecium salebrosum (Web. and Mohr) B. S. G. (1, 
2, 3, 5, 6, 7). Common in many habitats, especially on 
tree bases. 

Bryoerythrophyllum recurvirostrum (Hedw.) Chen (4). On 
thick humus, with Amblystegium juratzkanum, Myurella 
julacea and Platydictya jungermannoides. 

1974] Bryophytes — Stringer & Stringer 31 

Bryum angustirete Kindb. ex Mac. (1). On humus over 
sand, alone, and with Encalypta procera. 

Bryum argenteum Hedw. (1, 3). On dry humus and sand. 
Often with Funaria hygrometrica. 

Bryum creberrimum Tayl. (3, 5). On humus mixed with 
sand. Often with Ceratodon purpureus. 

Bryum pallescens Schleich. ex Schwaegr. (I, 2,3). Oh 
humus in moist habitats. 

Bryum  pseudotriquetrum (Hedw.) Gaertn. Meyer and 
Scherb. (2, 4). On thick, moist humus with Campylium 
stellatum and Brachythecium rutabulum. 

Callicladium haldanianum (Grev.) Crum (5, 6, 7). On 
humus and moist rotten wood. Previously reported only 
in the Winnipeg and Spruce Woods areas by Bird (1969) 
and by Stringer & Stringer (a, in preparation). 

Calliergon giganteum (Schimp.) Kindb. (4). On water- 
logged litter, in large masses mixed with Campylium 

Campylium chrysophyllum (Brid.) J. Lange (4). On well- 
rotted wood and humus. 

Campylium hispidulum (Brid.) Mitt. (4). One collection, 
on humus. with Amblystegium juratzkanum. 

Campylium stellatum (Hedw.) C. Jens. (4, 6). On wet 
humus or litter, often with Mnium affine or Thuidium 
delicatulum var. radicans. 

Ceratodon purpureus (Hedw.) Brid. (1, 3, 4, 5, 7). On 
humus over dry sand, with Bryum spp., Funaria hygro- 
metrica, or Polytrichum juniperinum. 

Dicranum bonjeanii De Not. ex Lisa (7). On thick humus 
over rotten wood, with D. polysetum and Ptilidium cili- 

Dicranum drummondii C. Müll. (6, 7). In thick, deep, pure 
clumps on humus, or as small plants on rotten wood with 
such typical species of decayed wood as Pohlia nutans 
and Tetraphis pellucida. 

Dicranum fuscescens Turn. (5, 7). On humus or rotten 
wood, usually in pure tufts. 

32 Rhodora [Vol. 76 

Dicranun polysetum Sw. (4, 5, 6, 7). Common, especially 
in jack pine forests of the area. Forms deep pure tufts. 
Often found also with Pleurozium schreberi. 

Distichium inclinatum (Hedw.) B. S. G. (4). Common in 
this collecting area and most often found with Myurella 
julacea, Amblystegium juratzkanum, and Platydictya 
jungermannoides. Several of the collections had mature 
capsules, and the large warty spores (354-404) distin- 
guish this species from the closely related Distichium 
capillaceum (Grout, 1936). 

Drepanocladus aduncus (Hedw.) Warnst. (1). On wet lit- 
ter under willows. 

Drepanocladus aduncus (Hedw.) Warnst. var. polycarpus 

. (Bland. ex Voit) Roth (1, 2). More commonly found 

. than the preceding, this variety occurred on moist humus 
or litter, often with Leptodictyum riparium. 

Drepanocladus revolvens (Sw.) Warnst. (4). Several col- 
lections made on wet litter or humus, in pure mats or 
with Pleurozium schreberi and Thuidium recognitum. 

Drepanocladus uncinatus (Hedw.) Warnst. (4, 5, 6, 7). On 
rotten wood or humus, in drier habitats than other mem- 
bers of the genus. With Brachythecium spp. and Bryum 

Encalypta procera Bruch (1). One collection, on thin hu- 
mus over dry sand. 

Eurhynchium pulchellum (Hedw.) Jenn. (4, 6, 7). On 
thick humus or litter in both wet and dry habitats. 

Fissidens adianthoides Hedw. (4). One small collection, 
with Campylium stellatum, on loose, moist humus. Not 
previously recorded for the province. 

Funaria hygrometrica Hedw. (1, 4, 5, 7). Frequent on 
sandy soil, often mixed with ashes of old fires. Com- 
monly with Bryum spp., Ceratodon purpureus, and Lepto- 
bryum pyriforme. 

Haplocladium microphyllum (Hedw.) Broth. (2, 3, 6, 7). 
Almost all collections on rotten wood but one collection 
(7) on humus. 

1974] Bryophytes — Stringer & Stringer 33 

Hedwigia ciliata (Hedw.) P. Beauv. (7). Several collec« 
tions, all on humus over rock or loose conifer needles over 

Hylocomium splendens (Hedw.) B. S. G. (4, 6, 7). On litter 
and loose humus, often with Pleurozium schreberi. 

Hypnum lindbergii Mitt. (4, 5, 6). Usually on wet, well- 
rotted wood or wet humus. 

Hypnum pallescens (Hedw.) P. Beauv. (3, 5, 6, 7). Com- 
mon on live and fallen bark of both conifers and hárd- 
woods. Often associated with Pylaisiella polyantha. 

Hypnum pratense Koch ex Spruce (1). One collection, on 
wet rotten bark with Drepanocladus aduncus var. poly- 

Isopterygium turfaceum (Lindb.) Lindb. (7). One collec- 
tion, on thick humus over granite rock, with Pohlia nu- 
tans and Brachythecium rutabulum. 

Leptobryum pyriforme (Hedw.) Wils. (1, 2, 4, 7). Com- 
mon on a wide variety of substrates and with a wide 
range of associated species. 

Leptodictyum riparium (Hedw.) Warnst. (1, 2). On wet 
humus or litter, often with Drepanocladus aduncus or 
Amblystegium varium. 

Leptodictyum trichopodium (Schultz) Warnst. var. kochii 
(B. S. G.) Broth. (1, 2). On deadfall or humus in moist 

Leskea polycarpa Hedw. (1, 2,5). On bark or humus, found 
most often with Amblystegium varium or Leptodictyum 
trichopodium var. kochii. 

Mnium affine Bland. ex Funck (4, 6). On wet humus or 
disintegrated rotten wood. 

Mnium cuspidatum Hedw. (1, 2, 3, 5, 6). On humus, rotten 
wood or litter in drier habitats than M. affine, often alone, 
or with Brachythecium salebrosum. 

Myurella julacea (Schwaegr.) B. S. G. (4). Common in 
wet habitats in this collecting area and often found with 
Campylium stellatum and Aulacomnium palustre. 

Orthotrichum obtusifolium Brid. (3). One collection, on 
poplar bark, with Hypnum pallescens. 

34 Rhodora [Vol. 76 

Orthotrichum speciosum Nees ex Sturm (3, 7). On live 
poplar bark, often with Pylaisiella polyantha, but per- 
sisting on deadfall even in advanced stages of decay 
where it often occurs with Haplocladium microphyllum. 

Platydictya jungermannioides (Brid.) Crum (4). Fairly 
common in this collecting area, with Myuwrella and Dis- 
tichium. Recorded only once previously for the province, 
as Amblystegiella sprucei (Bruch) Loeske, by Crum and 
Schofield (1959). 

Pleurozium schreberi (Brid.) Mitt. (3, 4, 5, 6, 7). Com- 
mon, and often forming large pure colonies in both wet 
and dry coniferous forests. 

Pohlia nutans (Hedw.) Lindb. (5, 6, 7). Usually on well- 
rotted wood but sometimes (7) on humus. 

Polytrichum juniperinum Hedw. (5, 6, 7). Usually on hu- 
mus over sand in drier habitats. 

Polytrichum piliferum Hedw. (7). One collection on humus 
over sand. 

Ptilium crista-castrensis (Hedw.) De Not. (5, 7). Usually 
a forest floor species of dry coniferous forests in the 
area, occurring on loose humus or needle litter over sand. 

Pylaisiella polyantha (Hedw.) Grout (2, 3, 5, 6, 7). Com- 
mon with Orthotrichum speciosum, especially on bark of 
live hardwoods, but also on humus (6, 7) or at the base 
of jack pine (5). 

Tetraphis pellucida Hedw. (6). On well-rotted wood, with 
Pohlia nutans and Drepanocladus uncinatus. 

Thuidium delicatulum (Hedw.) B. S. G. var. radicans 
Crum, Steere & Anderson (4). On loose wet humus and 
litter with Hypnum lindbergii, Campylium stellatum and 
Tomenthypnum nitens. 

Thuidium recognitum (Hedw.) Lindb. (4, 6). On loose hu- 
mus and needle litter in wet or moist habitats. 

Tomenthypnum nitens (Hedw.) Loeske (4). On loose, wet 
humus and litter. 

Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb. (7). On 
thin humus over sand. 

Stringer & Stringer 35 

1974] Bryophytes 


A search of the literature reveals that little work has 
been done on the bryophytes of Manitoba. Bird (1966), 
who gives a comprehensive list of the recordings for the 
province, lists only 289 taxa of bryophytes for the whole of 
Manitoba. This list has been somewhat augmented by Bird 
(1969) and Longton (1972), but published work on the 
bryophyte flora of the province is still greatly lacking. This 
situation is especially unfortunate as Manitoba is particu- 
larly interesting from a phytogeographic point of view, ly- 
ing as it does at a “crossroads of floral elements, from both 
north and south, east and west" (Lóve, 1959). 

Approximately half of the species recorded in the present 
study are pan-North American in distribution, according to 
the designations given by Bird (1969) and Bird and Won 
Shic Hong (1969). However, the Boreal element is repre- 
sented by species such as Dicranum fuscescens, D. polyse- 
twm, Tetraphis pellucida, Myurella julacea, Thuidium deli- 
catulum var. radicans, Thuidium recognitum, Pleurozium 
schreberi, Ptilium crista-castrensis, and Pylaisiella poly- 
antha. The Arctic Boreal element is represented by Di- 
cranum | bonjeanüi, Bryoerythrophyllum | recurvirostrum, 
Bryum pallescens, Orthotrichwm speciosum, Calliergon gi- 
ganteum, Campylium stellatum, Eurhynchium pulchellum, 
Tomenthypnum nitens, Hylocomium splendens, Polytrichum 
juniperinum, and Polytrichum piliferum. A few Eastern 
Boreal species are also present, i.e. Dicranum drummondii, 
Leptodictyum trichopodium var. kochii and Callicladium 

The hepatics are largely pan-North American species. 
Cephaloziella rubella, however, is Boreal and Ptilidium 
ciliare is Arctic Boreal (Bird & Won Shic Hong, 1969). 
Calypogeia . muelleriana and Cephalozia media have been 
previously recorded only from the north-east part of the 
province, from Churchill and York Factory respectively 
(Longton, 1972). The only previous record of Cephaloziella 
rubella was from Spruce Woods Provincial Park ( Stringer 

36 Rhodora [Vol. 76 

& Stringer, in preparation). Lophocolea minor, recorded 
from Grand Beach as new to the province, has since been 
found at several stations in Birds Hill and Whiteshell Pro- 
vineial Parks, where Radula complanata is also quite fre- 
quent. Riccardia pinguis has only two previous records, 
both from Spruce Woods Provincial Park, although subse- 
quent work by the authors shows that it is common on the 
calcareous clays of the southern part of the province. All 
of these liverworts are probably much more widely dis- 
tributed throughout the province than the scant records 

Several moss species which, according to the literature, 
should have a wide distribution have few records in Mani- 
toba. This is especially true of Brachythecium spp., per- 
haps because of the apparent scarcity of fertile material 
and the consequent difficulty in making a positive identifi- 
cation. Brachythecium rutabulum, common in the Grand 
Beach aréa studied, was reported as new to the province 
by Longton (1972), although Mueller-Dombois (1964) de- 
scribed it as one of the mosses forming the typically con- 
tinuous carpet in one of the jack pine — black spruce forest 
types in southeastern Manitoba. The only other records of 
Brachythecium campestre are also from the southern part 
of the province (Stringer & Stringer, in preparation, 1973) 
although this species appears to have a pan-North Ameri- 
can distribution. 

Callicladium haldanianum, reported for the first time in 
the province by Bird (1969) as Heterophyllium haldani- 
anum (Grev.) Kindb., reaches its western limit in Mani- 
toba. The Grand Beach record is further north than either 
of the Bird records. 

Fissidens adianthoides, a new record for Manitoba, ap- 
pears to be rare in western Canada (Bird 1966), although 
Grout (1936) gives its distribution as from Cape Breton 
Island to Vancouver Island and south to California and 

It is intended that the information presented in this 
study should, in conjunction with information presented in 

1974] Bryophytes — Stringer & Stringer 37 

further papers in this series, assist in describing the bryo- 
phyte flora of Manitoba and its relationships with the floras 
of adjacent regions. 


The field work was supported in part by Grant No. 140- 
118 from the University of Winnipeg. The authors also 
wish to express their gratitude to Dr. Dale H. Vitt, Univer- 
sity of Alberta, for his prompt assistance in identification 
of some of the Brachythecium material. 


Birp, C. D. 1966. A catalogue of the bryophytes reported from 
Alberta, Saskatchewan and Manitoba. Processed publication, 
University of Calgary, Calgary, Alberta. 

. 1969. Bryophytes of the aspen parkland of west- 
central Canada. Can. Jour. Bot. 47: 187-212. 

. & Won Suic Hone. 1969. Hepaticae of southwestern 
Alberta, Can. Jour. Bot. 47: 1727-1746. 

Crum, H. 1971. Nomenclatural changes in the Musci. The Bryol- 
ogist 74: 165-174. 

. & W. B. SCHOFIELD. 1959. The mosses of Gillam, Mani- 
toba. Bull. No. 160, Nat. Mus. Can., Ottawa, 

.„ W. C. STEERE & L. E. ANDERSON. 1965. A list of the 
mosses of North America. The Bryologist 68: 377-434. 

Grout, A. J. 1936. Moss flora of North America. Vol. I, Part I. 
Newfane, Vermont. Published by the author. 

LoNGTON, R. E. 1972. Nineteen bryophytes new to Manitoba. The 
Bryologist 75: 348-350. 

LóvE, D. 1959. The postglacial development of the flora of Mani- 
toba: a discussion. Can. Jour. Bot. 37: 547-585. 

MUELLER-DoMBors, D. 1964. The forest habitat types in south- 
eastern Manitoba and their application to forest management. 
Can. Jour. Bot. 42: 1417-1444. 

RowE, J. S. 1959. The forest regions of Canada. Bull. 125, Dept. 
Northern Affairs and Nat. Resources, Forestry Branch. Ottawa. 

SAYRE, G., C. E. B. BONNER, & W. L. CULBERSON. 1964. The authori- 
ties for the epithets of mosses, hepatics, and liehens. The Bry- 
ologist 67: 113-135. 

ScHUSTER, R. M. 1969. The Hepaticae and Anthocerotae of North 
America east of the hundredth meridian. Vol. II. 1062 pp. Colum- 
bia University Press, New York. 

38 Rhodora [Vol. 76 

1953. Boreal Hepaticae, a manual of the liver- 
worts of Minnesota and adjacent regions. Amer. Midland. Nat. 
49(2) : i-v, 257-684, 

SCOGGAN, H. J. 1957. Flora of Manitoba. Bull. 140, Biol. Ser. No. 
47, Nat. Mus. Can., Ottawa. 

STRINGER, M. H. L. & P. W. STRINGER. Studies on the bryophytes 
of southern Manitoba. I. Collections from Spruce Woods Pro- 
vincial Park, In preparation. 

& . 1973. Studies on the bryo- 
phytes of southern Manitoba. II. Collections from the Winnepeg 
area. Can. Field Nat. 87: 141-144. 






During the course of recent botanical studies in north- 
western Saskatchewan, several new or otherwise interest- 
ing plant records were collected. Most of these were col- 
lected from along the Green Lake-La Loche Road during a 
study supported by the Institute for Northern Studies, Uni- 
versity of Saskatchewan, or from the Lake Athabasca area. 
The purpose of this paper is to comment on these records 
in order to make this information generally available to 
taxonomists and phytogeographers. The collection numbers 
are those of the senior author unless otherwise indicated. 
All specimens cited have been deposited in the Fraser Her- 
barium, University of Saskatchewan (SASK). 

SPARGANIUM CHLOROCARPUM Rydb. 7 mi. s.-s.e. of La 
Loche, Mile 60 Buffalo Narrows-La Loche Road, Harms 
17635; McAneely Creek, 2 mi. s.w. of Turnor Lake village, 
Harms 18390; Bear Creek, Mile 41 Buffalo Narrows-La 
Loche Road, Harms 19661. 

This species has never been listed for Saskatchewan by 
Fraser and Russell (1937), Fraser and Russel] (revised by 
Russell, Ledingham and Coupland (1954)), or Breitung 
(1957). The specimens of the species were for the most 
part confused with Sparganium multipedunculatum. While 
a taxonomic problem exists between the two taxa, and even- 
ually both of them, together with S. angustifolium, may be 
merged as varieties under an enlarged S. simplex, there is 
no question that many of our specimens best fit S. chloro- 
carpum as it is now recognized. Other specimens in the 
Fraser Herbarium now identified as S. chlorocarpum in- 
clude the following: 

Melfort, J. B. Millar 64-163, 64-12; Saskatoon, J. H. 
Hudson 2488, 2548, J. B. Millar 65-217, W. P. Fraser 1930; 


40 Rhodora [Vol. 76 

Pike Lake, 20 mi. s. of Saskatoon; W. P. Fraser 1 Sept. 
1937, R. C. Russell, et al., 4 August 1937; St. Denis, J. B. 
Millar 62-72; W. of Whitefox, W. P. Fraser & G. F. Leding- 
ham 28 July 1936; Swift Current, J. L. Bolton 8 July 1936; 
Dahlton, W. P. Fraser and. G. F. Ledingham 20 July 1937; 
Egg Creek, s. of Cumberland House, G. H. Townsend 55; 
Waskesiu, W. P. Fraser 24 July 1938; Lake Waskesiu Road, 
W. P. Fraser July 1932; Mile 23 Waskesiu Road to Prince 
Albert, W. P. Fraser 24 August 1934; Meadow Lake Pro- 
vincial Park, Waterhen River, J. H. Hudson 2427; Sylvan 
Lake, Candle Lake Road, J. K. Jeglum 18 July 1964; 1.7 mi. 
n. of forks at Candle Lake, J. K. Jeglum 9 July 1964. 

CALAMAGROSTIS LAPPONICA (Wahl.) Hartm. 3 mi. s. of 
La Loche, Mile 64 Buffalo Narrows-La Loche Road, Harms 
17532, 17565; 2 mi. s. of La Loche, Mile 65 Buffalo Nar- 
rows-La Loche Road, Harms 17547, 17543; Island in Wil- 
liam River, south shore Lake Athabasca, R. Hermesh 485; 
Thompson Bay, south shore of Lake Athabasca, R. Hermesh 

This northern species has never been previously reported 
for Saskatchewan. There seems to be intergradation be- 
tween Calamagrostis lapponica and C. neglecta (Ehrh.) 
Gaertn. as indicated by an intermediate collection from 
Taylor Lake, Mile 19 Buffalo Narrows-LaLoche Road 
(18160), and between C. lapponica and both C. neglecta 
and C. inexpansa A. Gray as indicated by intermediate 
specimens from the same area (18142, 18147). The whole 
C. lapponica, C. neglecta, and C. inexpansa complex appears 
in need of careful taxonomic study. 

CAREX ADUSTA Boott. Turnor Lake, Harms 18364-B, 

In the Fraser Herbarium, there are three specimens from 
the Waskesiu area or the road to Montreal Lake, Prince 
Albert National Park (W. P. Fraser July 8, 1939, June 25, 
1940, June 22, 1941). This species has been collected by 
J. H. Hudson at Flin Flon, Manitoba, near the Saskatche- 
wan border. Breitung (1957) reports it also from Meadow 

1974] Saskatchewan Plants — Harms & Hudson 41 

Lake but a substantiating voucher is missing from the 
Fraser Herbarium. The above record represents a con- 
siderable northwestward range extension in Saskatchewan. 

CAREX RUFINA Drej. Beach at Thomson Bay, south shore 
Lake Athabasca, R. Hermesh 421. 

This represents the first report for this species in Sas- 
katchewan. Characteristic material of this species comes 
from Greenland and Iceland. Scoggan (1957, p. 179) re- 
lates that A. E. Porsild collected typical material from 
southern Keewatin, Northwest Territories, while Baldwin 
found abberrant material at Nueltin Lake, northern Mani- 
toba, about 125 miles south of Porsild's station. Porsild 
has annotated Baldwin’s sheet as follows: “Differs from 
typical C. rufina by paler scales having more prominent 
midveins, paler and more prominently nerved perigynia, 
and by terminal spikelets being not so strictly gynaecan- 
drous (in some even androgynous). Perigynia only rarely 
with a few teeth in the upper part." The present Lake 
Athabasca material agrees well with the characters men- 
tioned in these statements. This collection represents about 
a 350 mile westward range extension for the species. 

HABENARIA ORBICULATA (Pursh) Torr. Lac La Plonge, 5 mi. 
e. of Beauval, Harms 17959-B. 

In the Fraser Herbarium there are specimens of this 
species from Big Sandy Lake, Hansen Lake Road (G. W. 
Argus 4237), from La Ronge (J. S. Maini 611), and from 
Pinkney Lake, 35 mi. n.e. of Candle Lake (J. M. A. Swan 
65-45). Breitung (1957) also reports it from Torch River 
and Amisk Lake, the latter based on a collection by J. H. 
Hudson. The present record is the first one from the west- 
ern part of the province and represents a considerable west- 
ward extension of the main species range. It is disjunctly 
present in British Columbia. 

SPIRANTHES GRACILIS (Bigel) Beck. (S. lacera Raf.). 
Little Amyot Lake area, Mile 69 Green Lake-Buffalo Nar- 
rows Road, Harms 17736; 2 mi. e. of Beauval, Harms 

42 Rhodora [Vol. 76 

17776; Taylor Lake, Mile 19 Buffalo Narrows-La Loche 
Road, Harms 18139-B. 

There are specimens of this species in the Fraser Her- 
barium from Lake Waskesiu, Prince Albert National Park 
(Anonymous July 1932) and from 20 miles south of Mea- 
dow Lake (A. J. Breitung 8164). The above records am- 
plify the known range of the species and extend it to north- 
western Saskatchewan. 

SPERGULA ARVENSIS L. 2 mi. w. of Beauval, Harms 17846. 

Breitung (1957) does not report this plant for Saskatche- 
wan, and Scoggan (1957) does not report it for Manitoba. 
Boivin (1968) listed this species as doubtful for Saskatche- 
wan. Hudson (1972) reported it as new to Saskatchewan 
on the basis of a collection from Gronlid (B. Zuk 1971, 
DAO). Boivin, in personal correspondence to J. H. Hudson, 
indicated the presence of an earlier collection from Sylvania 
(K. Drake Juiy 26, 1966, DAS). Mrs. H. D. Bobier recently 
sent a collection of the species from Rapid View (west of 
Meadow Lake) to J. H. Hudson. A map by J. F. Alex indi- 
cates two infestations of this weed near Outlook, Saskatche- 
wan but these are not substantiated by voucher specimens. 
Thus the above record represents only the fourth report for 
Saskatchewan and the only one from this far northwest in 
the province. 

CALTHA NATANS Pallas. 7 mi. s.-s.e. of La Loche, Mile 60 
Buffalo Narrows-La Loche Road, Harms 17633. 

There are no Saskatchewan specimens of this species in 
the Fraser Herbarium, Breitung (1957) reported it from 
Windrum Lake (56°02’N, 104°W) and Amisk Lake, the 
latter based on a collection by J. H. Hudson. The present 
collection presumably represents the third report for Sas- 

RANUNCULUS HYPERBOREUS Rottb. Open beach, William's 
Point, south shore Lake Athabasca, R. Hermesh 415. 

This represents the first record of this northern butter- 
cup species in Saskatchewan. 

1974] Saskatchewan Plants — Harms & Hudson 43 

Mile 70 Green Lake-Buffalo Narrows Road. 

Breitung (1957) omits this species from his list of Sas- 
katchewan plants. Boivin (1968) reports only 3 collections 
from the prairie provinces: Cochrane River, Reindeer 
Lake, and Lake Axis. The present collection would there- 
fore represent only the second Saskatchewan record for the 

EUPHRASIA ARCTICA Lane var. DOLOSA Boivin (E. subarctica 
Raup, E. disjuncta Fern. & Wieg. var. dolosa Boivin). 2 mi. 
s. of La Loche, Mile 65 Buffalo Narrows-La Loche Road, 
Harms 17550. 

In the Fraser Herbarium, the only Saskatchewan speci- 
men of this species is from Stony Rapids, north of Fond-du- 
lac River (J. S. Maini & M. Swan Aug. 3, 1961). Breitung 
(1957) cites a Raup report from Lake Athabasca. The 
above record represents about a 200 mile southward range 
extension for the species in Saskatchewan. 

LOBELIA DORTMANNA L. Little Amyot Lake, Mile 70 Green 
Lake-La Loche Road, Harms 16972. 

In the Fraser Herbarium we have specimens of this spe- 
cies from Little Gull Lake, south shore Lake Athabasca 
(G. W. Argus 553-63) and Carswell Lake, 40 mi. south of 
Lake Athabasca (G. W. Argus 628-62). Breitung (1957) 
reports the species also from Windrum Lake, north of the 
Churchill River, 56°02’N, 104°W. Therefore, the above rec- 
ord represents the third general locality for the species in 

ADOXA MOSCHATELLINA L. 17 mi. n. of Meadow Lake, J. H. 
Hudson 2729 (SASK). 

Boivin (1972, p. 5) reports this species in Saskatchewan 
from Pasquia Hills and Candle Lake; the latter specimen, 
which he collected, was the first record for Saskatchewan. 
The present material is evidently the third record of the 
species for the province and represents more than a 100 
mile westward range extension in the province. 

44 Rhodora [Vol. 76 

ANAPHALIS MARGARITACEA (L.) B. & H. 3 mi. s. of Buffalo 
Narrows, Mile 124 Green Lake-Buffalo Narrows Road, 
Harms 18244. 

There are various specimens of this species in the Fraser 
Herbarium from the Cypress Hills area of Saskachewan. 
Breitung (1957) reports it also from Cut Knife but a sub- 
stantiating voucher has not been found. Therefore the 
above record represents at least a 200 mile, if not a 300 
mile northward range extension. 

BIDENS BECKII Torr. Little Amyot Lake, Mile 70 Green 
Lake-Buffalo Narrows Road, Harms 16978. 

There are two specimens of this species in the Fraser 
Herbarium from the Cumberland House area on the Sas- 
katchewan River (A. E. Etter Aug. 8, 1934; D. Dabbs 126- 
66) in east-central Saskatchewan. The above report there- 
fore represents the second known Saskatchewan locality for 
the species and is a considerable westward range extension 
in the province. 


BorviN, B. 1968. Flora of the Prairie Provinces, Part II. Reprinted 
from Phytologia 16-18. 

1972. Flora of the Prairie Provinces, Part III. Re- 
printed from Phytologia 22-23. 

BREITUNG, A. J. 1957. Annotated Catalogue of the Vascular Flora 
of Saskatchewan. The American Midland Naturalist 58: 1-72. 

FRASER, W. P. & R. C. RUSSELL. 1937. An Annotated List of the 
Plants of Saskatchewan. University of Saskatchewan, Saska- 

HUDSON, J. H. 1972. 1971 Botanical Records for Saskatchewan. 
The Blue Jay 30: 122-124. 

vision of Fraser, W. P. and R. C. Russell, An Annotated List 
of the Plants of Saskatchewan. University of Saskatchewan, 

SCOGGAN, H. J. 1957. Flora of Manitoba. National Museum of 
Canada Bulletin No. 140, Ottawa. 




When plants of a witchweed were discovered in 1956 
parasitizing Zea mays L. in North and South Carolina, 
specimens were sent to Kew for identification. I determined 
them as Striga asiatica (L.) O. Kuntze and communicated 
the name to Dr. S. F. Blake and other authorities in the 
United States, who used it until a paper by Dr. C. J. 
Saldanha (1963) proposed the rejection of that name in 
favour of S. lutea Lour. (Smith 1966). The purpose of this 
paper is to reinstate S. asiatica on the basis of typification 
of the Linnean basionym Buchnera asiatica, rather than 
on the opinions of other taxonomists. 

Linnaeus’ description of Buchnera asiatica in ‘Species 
Plantarum' (1753) is unusually full for that work and 
entirely devoid of any references to literature. This indi- 
cates that he had before him a specimen, or specimens, 
from which he drew up his description. Examination of 
his herbarium at the Linnean Society, London, revealed 
six sheets bearing the name ‘Buchnera asiatica! in Linnaeus’ 
hand and now numbered according to Savage's Catalogue 
as 790/10, 11, 12, 13, 14, & 15. 

Sheets 13, 14 and 15 can be discounted, since they are 
annotated as having been collected by Thunberg (13) some- 
time after 1772, and by Koenig (14 & 15) in 1777 according 
to the date on the sheet. They are referable to Striga 
bilabiata (Thunb.) O. Kuntze (syn. S. thunbergii Benth.) 
and S. euphrasioides Benth., respectively. 

Sheet 12 bears a single simple plant with pale flowers 
that would be known in India as Striga lutea. The only 
annotation on the sheet is ‘Indica,’ indicating its country 
of origin. The specimen does not fit Linnaeus' description, 
and it can be ruled out as the type. 

Five specimens are glued to sheet 10, including a well- 
branched plant with dark-coloured flowers that fits the 


16 Rhodora [Vol. 76 

original description admirably. Two other specimens match 
it except for their simple habit. The remaining two are 
only the inflorescences of Striga densiflora Benth. The 
specimen on sheet 11 is very similar to the large branched 
one on sheet 10, but there is no annotation apart from the 
name. On sheet 10, however, Linnaeus has added the 
locality “ins. Johan.," and although the ‘Species Plantarum’ 
cites the distribution as “Habitat in Zeylona, China” this 
reference to “ins. Johan.” provides a clear indication of the 
provenance and history of what must be regarded as the 
type material of Buchnera asiatica. Where, then, is “ins. 
Johan.” and who collected the specimen? 

Several voyages to China were undertaken before 1753. 
including one by Peter Osbeck who wrote an account of 
it in Swedish (1757) which was translated into German 
(1765) and from the German edition into English, and 
published in 1771 under the title ‘A voyage to China and the 
East Indies, together with a voyage to Suratte by Olof 
Toreen in a series of letters to Doctor Linnaeus.’ Toreen 
(or Torén), in fact, continued his voyage as far as China 
and it is his letters, and not Osbeck’s text, which are of 
particular interest since he tells (p.166) how, on 16 August 
1750, he landed at the island of “St. Joanna” in the Comoros 
near Madagascar. It is evident that Linnaeus worked out 
the collections made by Toreen and Osbeck, attributing 
them al] to China in the appendix to Osbeck and Toreen’s 
travels entitled ‘Flora Sinensis.’ There, on p. 356, is listed 
Buchnera asiatica with the reference to ‘Species Plantarum.’ 
It is interesting to note that the same island was visited by 
Koenig in 1768 where he gathered the plant Linnaeus 
(1771) described as Baccharis arborea (which Brenan 
(1968) identified as Vernonia grandis (DC.) Humbert var. 
comorensis Humbert), yet Linnaeus gave the occurrence as 
the “East Indies.” 

Striga asiatica (L.) O. Kuntze, Rev. Gen. Pl, 2: 466. 1891. 
Type. Comoro Islands, St. Joanna, Toreen LINN. (Sav- 
age Cat. No. 790/10 partly). 

1974] Striga — Hepper AT 

Buchnera asiatica L., Sp. Pl. 630. 1753. 

Striga lutea Lour., Fl. Cochinch. 22. 1740. 

Buchnera hirsuta Benth., Scroph. Ind. 41. 1835. 

Striga hirsuta, (Benth.) Benth. in Hook., Comp. Bot. Mag. 
1: 363. 1836. 

Bentham's work was completed long before an interna- 
tional code of botanical nomenclature had been agreed, and 
certainly his concepts of typification were not in accord 
with modern attitudes. Hence, he coined the new names 
to replace Buchnera asiatica which seemed to him, as to 
Saldanha, to apply to a number of different species. The 
more so since he considered S. lutea Lour. to be a distinct 

Saldanha considered Striga asiatica and its basionym as 
nomena rejicienda under Art. 69 since “any attempt to 
decide a lectotype is bound to be arbitrary and open to 
question." On the contrary, this species is one of the few 
Linnean species that may be typified and localized with 

I am grateful to my colleague Mr. R. D. Meikle for his 
helpful advice and assistance. 


BRENAN, J. P. M. 1968. The identity of Baccharis arborea L. 
Kew Bull. 21: 427. 

LiNNAEUS, C. 1753. Species Plantarum, 630. 

1771. Mantissa Plantarum, 284. 

OsBECK, P. 1771. A voyage to China and the East Indies. (English 

SALDANHA, C. J. 1963. The genus Striga Lour. in Western India. 
Bull. Bot. Surv. India 5 (1) : 67-70. 

SMITH, C. EARLE. 1966. Identity of witchweed in the southeastern 
United States. Rhodora 68 (774) : 167. 




The problematic taxon long-called Senecio rugelia (Shuttl. 
ex Chapman) A. Gray (Compositae) is a distinctive species 
occurring in coniferous associations along the higher ridges 
of the Great Smokies astride the North Carolina-Tennessee 
border. It is a tall, coarse herb with several conspicuous 
discoid heads and dull, dirty-white corollas. The species 
was introduced to science as Rugelia nudicaulis Shuttl. ex 
Chapman in 1860, but in 1883 Asa Gray transferred it to 
Senecio, where it has remained. 

Many botanists have recognized the incongruity of in- 
cluding this entity in Senecio, but the morphological toler- 
ances of Senecio have been usually treated as very wide, 
and so the matter has not been pursued. However, Green- 
man (1901) and Alexander (1937) noted the possible 
relationship to Cacalia. 

The species referable to Cacalia have had their systematic 
woes, being variously regarded as constituting several gen- 
era, or as members of the single genus Cacalia, or as a part 
of Senecio. Compounding these difficulties has been a dis- 
agreement concerning the typification of the name Cacalia. 
A review of these matters is beyond the scope of this paper, 
but we are treating Cacalia in the expanded sense, and we 
accept Cacalia hastata L. as the generic type (Vuilleumier 
& Wood, 1969). 

In general, Cacalia differs from Senecio by the two floral 
features of discoid heads and white to creamy corollas. 
Both are features which occur independently as evolution- 
ary tendencies in other alliances within Senecio. In Cacalia, 
however, the two occur together, presumably defining a 

natural assemblage. 

*Contribution no. 1193. Division of Biology, Kansas Agr. Expt. 
Station, Manhattan. Financial support from the National Science 
Foundation is gratefully acknowledged. 


1974] Caealia — Barkley and Cronquist 49 

Cacalia is here regarded as a derivative of Senecio section 
Palmatinervii of Mexico, a group of coarse herbs strongly 
resembling Cacalia in aspect. Nearly all species of the 
Palmatinervii for which chromosome counts are reported 
have n=30 (Gibson, 1969). Cacalia has n—30, and a re- 
duction series of n=28, 27, 26, 25, and 20 (Ornduff et al., 
1963, 1967). The species here offered as Cacalia rugelia 
has »—28, a number unreported for Senecio (Fedorov, 

In a recent study of Mexican cacalioids, Pippen (1968) 
recognizes four genera, but he offers a table summarizing 
the distinctions between these ‘“‘Cacalioid genera” and Sene- 
cio (table 2, p. 371). Incorporated into the table are the 
cacalioid features of discoid heads and white or creamy 
florets. The table also shows that the *Cacalioid genera" 
characteristically have deeply lobed disk corollas, a feature 
which we find in Cacalia rugelia as well. Furthermore, in 
Pippen's key the present species falls into the assemblage 
treated as the Mexican segregate-genus Odontotrichum, and 
it is morphologically compatible with that group. The 
floristic relationships between the Southern Appalachian 
region and some of the mountains of Mexico have been 
noted by other authors (e.g. Sharp, 1946). 

In the light of the foregoing considerations, the following 
transfer is proposed: 

Cacalia rugelia (Shuttl. ex Chapman) Barkley & Cronquist, 

comb. nov. 

Type: “Smoky Mountains, Tennessee. Rugel, Buckley." 
Paratype: Buckley, NY! 

Senecio rugelia A. Gray, Proc. Am. Acad. 19: 54. 1885. 

Rugelia nudicaulis Shuttleworth ex Chapman, FI. S. U. S. 
246. 1860. 
non Cacalia nudicaulis (Less.) O. Kuntze, 1891. 
non Senecio nudicaulis Buck.-Ham. ex D. Don, 1825. 

ALEXANDER, E. J. 1937. Senecio rugelia.  Addisonia 20: 29-30, 
pl. 655. 

50 Rhodora [Vol. 76 

FEDOROV, AN. A., ed. 1969. Chromosome numbers of flowering 
plants. Soviet Sciences Press. Leningrad. 

GIBSON, E. S. 1969. A revision of the section Palmatinervii of the 
genus Senecio (Compositae) and its allies. Unpub. doctoral dis- 
sertation, Kansas State Univ. Manhattan. pp. 1-178 + VI. 

GREENMAN, J. M. 1901. Monogr. nord-und centralam. Arten der 
Gattung Senecio. Leipzig. (Reprinted in Englers Bot. Jahrb. 32: 
1-33. 1902.) 

ORNDUFF, R. P. H. Raven, D. W. KvHos & A. R. KINCHEBERG. 
1963. Chromosome numbers in Compositae III. Senecioneae. 
Am. Jour. Bot. 50: 131-139. 

OnNDUFF, R., T. Mosquin, D. W. KvHos & P. H. RAVEN. 1967. 
Chromosome numbers in Compositae VI. Senecioneae II. Am. 
Jour. Bot. 54: 205-213. 

PIPPEN, R. W. 1968. Mexican “Cacalioid” genera allied to Senecio 
(Compositae). Contr. U. S. Natl. Herb. 34: 365-447. 

SHARP, A. J. 1946. Informe preliminar sobre alqunos estudios 
fitogeograficos efectuados en Mexico y Guatemala. Rev. Soc. 
Mexicana Hist. Nat. 7: 35-39. figs. 1-5. 

VUILLEUMIER, B. S. & C. E. Woop. 1969. Lectotypification of Caca- 
lia L. (Compositae-Senecioneae). Jour. Arnold Arb. 50: 268-275. 





The names Scirpus americanus Pers. (Syn. Pl. 1: 68. 
1805) and Scirpus pungens Vahl (Enum. Pl. 2: 255. 1805) 
are usually applied to the same species by Cyperologists 
(Beetle, 1947; Koyama, 1963). The former name for plants 
commonly known as Three-square in North America pres- 
ently receives widespread usage because of priority. Data 
from type specimens and original descriptions now make it 
apparent that these names should be applied to different 
species. The type of S. americanus (Hab. in Carolina in- 
feriore, Herb. Michx., P) is conspecific with plants usually 
treated (Beetle, 1947; Koyama, 1963) as S. olneyi Gray 
(Bost. Journ. Nat. Hist. 5: 238. 1845). The type of S. 
pungens (Erhart misit, Herb. Vahl, C) is conspecific with 
plants usually treated (op. cit.) as S. americanus. 

In his original description of Scirpus americanus, Per- 
soon cites Michaux's description of S. triqueter L. (Fl. Bor.- 
Am. 1: 30. 1803.) and gives the habitat as "in Carolina 
inferiore." The specimens on the sheet thus named and 
labeled in Michaux's herbarium at P have strongly winged 
culms, short involucral bracts, shallowly cleft scales, small 
achenes, and broad bristles characteristic of plants resem- 
bling the type of S. olneyi (Rhode Island, Providence, Olney, 
GH). Thus the correct name for plants usually treated as 
S. olneyi is S. americanus. 

In the original description of Scirpus pungens, Martin 
Vahl gave the habitat as, *in Europa, in insula Borboniae. 
Commerson." The specimen from Vahl's herbarium desig- 
nated as the type probably was collected in Europe and also 
contains the word “pungens Vahl" presumably in the hand- 
writing of Jens Vahl, son of Martin Vahl. No specimen 
labeled, *in insula Borboniae," or collected by Commerson 
has been located. The triangular culms, long involucral 


52 Rhodora [Vol. 76 

bracts, deeply cleft scales, and narrow bristles are char- 
acteristics of plants usually but incorrectly treated as S. 
americanus. Scirpus pungens is the earliest available name 
for these plants. 

The literature cited by Vahl is also in accordance with his 
description and the designated type specimen. The cited 
illustrations of Morrison (Pl. Hist. Univ. Oxon. s. 8, t. 10, 
f. 20. 1699.) and Plukenet (Algm. Bot. t. 40, f. 1. 1696.) 
closely resemble the type specimen of Scirpus pungens. 
Roth's descriptions of S. mucronatus L. (Tent. Fl. Germ. 
1: 28. 1788, and 2: 60. 1789.) and S. triqueter L. (Neue 
Beytr. 1: 91. 1802.) cited by Vahl best apply to plants 
resembling the type specimen of S. pungens. Apparently 
Roth was confused about the identity of S. mucronatus and 
S. triqueter, and incorrectly applied the names to plants of 
S. pungens. 


I am grateful to H. Heine (P), J. Raynal (P), and A. 
Skovsted (C) for courtesies enabling me to examine critical 
specimens and data at their respective institutions, and to 
John Braxton (PH) and Helena Greenwood (PH) for their 
assistance. Abbreviations of institutions are those given 
by Lanjouw and Stafleu (1964). 


BEETLE, A. A. 1947. Scirpus. N. Am. Fl. 18: 481-504. 

KovAMA, T. 1963. The genus Scirpus Linn., critical species of the 
section Pterolepis. Canad. Jour. Bot. 41: 1107-1131. 

LANJOUW, J. and F. A. STAFLEU. 1964. Index Herbariorum. Part 1. 
The herbaria of the world, International Association for Plant 
Taxonomy, Utrecht. 251 pp. 




The most recent entry in the literature of Cardamine 
clematitis appears on page 508 of the Manual of the Vas- 
cular Flora of the Carolinas by Radford, Ahles and Bell 
(1968). They cite “Shuttlew.” as the authority for this 
taxon. This authority refers to Robert James Shuttle- 
worth, a famed naturalist and collector who was the spon- 
sor of Ferdinand Rugel, a prolific plant collector in eastern 
North America from 1840 and for many years thereafter 
until his death in 1879. 

The various references in the literature to Cardamine 
clematitis have consistently overlooked the fact that Shut- 
tleworth cannot be cited as the sole publishing authority for 
the specific epithet, clematitis. In addition to Radford, 
Ahles and Bell (1968), several other earlier North Ameri- 
can references also provided descriptions and occasionally 
illustrations of C. clematitis, and all accepted Shuttleworth 
as the sole publishing authority. These include: Chapman, 
Flora of the Southeastern United States, ed. 2, Supplement 
p. 605. 1887; ibid., ed. 3, p. 25. 1897; Britton & Brown, 
Illust. Flora of the Northern United States and Canada, ed. 
1. 2: 130. fig. 1730. 1897; ibid., ed. 2. 2: 185. fig. 2088. 
1913; Small, Flora of the Southeastern United. States, ed. 1, 
p. 482. 1903; ibid., ed. 2, p. 568. 1933; Fernald in Gray's 
Manual of Botany, p. 721. 1950; Gleason and Cronquist in 
Britton and Brown, Illust. Flora of the Northeastern United 
States and Adjacent Canada 2: 230, fig. p. 231. 1952; 
Gleason & Cronquist, Manual of Vascular Plants of North- 
eastern United States and Adjacent Canada, p. 340. 1963; 
and Radford, Ahles and Bell, Guide to the Vascular Flora 
of the Carolinas, p. 173. 1964. 

‘Research Botanist, Herbarium, U.S. National Arboretum, Agricul- 

tural Research Service, Northeastern Region, U.S. Department of 
Agriculture, Washington, D.C. 20002. 


54 Rhodora [Vol. 76 

In the course of evaluating the Ferdinand Rugel collec- 
tions in the Isaac C. Martindale Herbarium at the U.S. 
National Arboretum, F. G. Meyer called my attention to an 
unidentified specimen of Cardamine collected by Rugel and 
accompanied by a signed holograph label that reads “Top 
of Smoky Mts. on the first Branch Nord Carolina Side. 
May 1866." Although this Rugel specimen clearly repre- 
sents C. clematitis, it is not a part of the type. 

The first published reference to Cardamine clematitis is 
on page 53 of Sereno Watson's Bibliographic Index to 
North American Botany (Smithsonian Miscellaneous Col- 
lections No, 258, 1878) that reads as follows: “C. clematitis 
Shuttl. in Herb. Gray, ined." It is evident that Watson took 
the name from Shuttleworth’s printed herbarium exsiccata 
label which reads “Cardamine clematitis Shuttle. n. sp.” 
and that name was used only as a nomen nudum. Watson’s 
reference does not incorporate a validating description. On 
this basis alone, clematitis cannot be used as a valid specific 
epithet. It is listed, however, in Index Kewensis 1: 421. 
1895 as: “Clematitis Shuttle. ex S. Watson" ! 

Later, in a paper by Asa Gray entitled “Some New 
North American Genera, Species, &c." (Proc, Am. Acad. 
Arts Sci. n.s., 7:45. 1880, a valid Latin description is pro- 
vided. Gray precisely identified the plant as “Cardamine 
clematitis Shuttleworth in coll. distrib. Rugel," and explains 
that the original collection was from “wet ground along 
streamlets in the higher Iron or Smoky Mountains of North 
Carolina and Tennessee, collected in 1844 by Rugel .. .” 

When dealing with nomenclatural problems of this na- 
ture, we are guided by Article 46 and its accompanying 
recommendations (particularly Recommendation 46C) in 
the International Code of Botanical Nomenclature, p. 46. 
1972. 'The correct authority citation for Cardamine clema- 
titis is C. clematitis Shuttleworth ex A. Gray, although it 
may be shortened, particularly in floristic treatments, to 
cite only the publishing author. Under no circumstances 
can Shuttleworth be assigned solitary authorship. The cor- 
rect citation of the name is: 

1974] Cardamine clematitis — Dudley 55 

Cardamine clematitis Shuttleworth ex A. Gray, Proc. 

Amer. Acad. Arts Sci, new series, 7: 45. 1880 — non 

Shuttl. ex Watson, Bibliog. Index North American Bot., 

p. 53, 1878. 

The original collection and designated type of Cardamine 
clematitis is Rugel No. 19 collected in 1844 in the Smoky 
Mts. of Tennessee. The holotype is deposited at GH, and 
cited duplicates (isotypes) are to be found at G-Herb. Bois- 
sier and W. 

TYPE: Tennessee, Smoky Mountains [‘‘in locis humidis 
et ad regionis super. montium Smoky Ms., Tennessee, Mai 
1884.”] May 1844, Ferdinand Rugel No. 19 (holotype, GH; 
isotypes BM, G, W). 

The most complete set of Rugel’s material, purchased from 
Shuttleworth, is at the British Museum; however, reference 
to the existence of such a specimen has not previously ap- 
peared in the literature. N. K. B. Robson of the British 
Museum (Natural History) assures me that a duplicate is 
retained at BM. 

The one Rugel collection of Cardamine clematitis not 
previously reported in the literature is a specimen collected, 
probably a unicate, after Rugel’s professional contacts with 
Shuttleworth had ceased. The specimen was maintained in 
Rugel’s personal herbarium until that herbarium was pur- 
chased in 1881 from Rugel’s heirs by Isaac C. Martindale: 
North Carolina: “Top of Smoky Mts. on first Branch, Nord 
Carolina Side", Ferdinand Rugel, May 1866 (NA — from 
herbarium of Isaac C. Martindale). 

The astute monographer of Cardamine and indefatigable 
expert on the Cruciferae, D. E. Schulz, in Monographie der 
Gattung Cardamine” (Bot. Jahrb. 32: 440. 1903.) recog- 
nized Shuttleworth as responsible for the epithet clematitis, 
but also recorded that Asa Gray provided a description of 
the species in Proc. Am. Acad. Arts Sci. n.s. 7: 45. 1880. 
Schulz examined the original 1844 material of C. clematitis 
collected by Rugel (GH, G & w — but not BM), which was 
annotated and distributed by Shuttleworth; he also cited 
numerous additional collections made by J. K. Small, N. L. 

56 Rhodora [Vol. 76 

& E. G. Britton & A. M. Vail, S. B. Buckley, J. K. Small & 
A. A. Heller, W. M. Canby, M. E. Hyams and J. W. Chicker- 



A point should also be discussed that concerns the synon- 
omy of Cardamine clematitis Shuttleworth ex A. Gray as 
presented by the Radford, Ahles and Bell publications 
(1964 & 1968). These works refer “C. flagellaris” (= C. 
flagellifera Schulz, Bot. Jahrb. 32: 405. 1903.) into synon- 
omy under C. clematitis. Although the type specimen of 
C. flagellifera (Biltmore Herbarium 7756) was originally 
annotated as C. clematitis, it is not conspecific with C. 
clematitis. Not only does C. flagellifera stand morphologi- 
cally and ecologically distinct from C. clematitis, but it was 
also assigned by Schulz within Cardamine to Sect. Macro- 
phyllum, whereas C. clematitis was referred by Schulz to 
the typical section. Small (1903) described C. hugeri that 
Radford, Ahles & Bell (1964 and 1968) sank under C. 
clematitis. However, if referrable into synonomy at all, 
C. hugeri, a low altitude plant, more logically pertains to 
C. flagellifera. 

In Watson’s reference (loc. cit.) to Cardamine clematitis, 
he cites as a synonym a “Nasturtium officinale” that ap- 
peared in the Supplement to Torrey and Gray, A Flora of 
North America 1: 666. 1843. In addition to validating, 
describing and typifying C. clematitis, Gray (loc. cit.) 
explains Watson’s confusion in incorrectly citing this 
“Nasturtium officinale” as a synonym of C. clematitis. The 
original Rugel material, annotated and distributed from 
Switzerland by Shuttleworth, was a mixed collection. A 
part representing C. clematitis “was mixed up with a Flor- 
ida species intermediate between Cardamine and Nastur- 
tium, first received from Leavenworth without fruit, and 
referred in the supplement to the first volume of Torrey and 
Gray's Flora to N. officinale.” This element of “Nasturtium 
officinale”, according to Gray, was later received from S. B. 

1974] Digitaria sanguinalis — Ebinger 57 

Buckley; again later received from Shuttleworth’s dis- 
tributed collections of Rugel as Cardamine curvisiliqua 
Shuttleworth; and yet again received from Shuttleworth as 
Nasturtium stylosum Shuttleworth! The current fate of 
Leavenworth’s “Nasturtium officinale’ is not within the 
scope of this paper; however, both Chapman (1887) and 
Small (1903) equate Leavenworth’s “Nasturtium officinale” 
with Cardamine curvisiliqua Shuttleworth. 


a result of extensive field work in east-central Illinois an 
unusual form of the common crabgrass was found. This 
form differs from typical Digitaria sanguinalis (L.) Scop. 
in that long, spreading, papillose-based hairs are found 
scattered along both sides of the flattened rachis. These 
colorless hairs are 3-6 mm long (rarely 1 cm long) and on 
the specimens examined usually 1 to 5 hairs are found on 
each cm of rachis length. As a result of this difference the 
following form is described. 

Digitaria sanguinalis (L.) Scop. forma illinoensis Ebinger 
forma nova. 

A forma sanguinalis differt pilis base papillosis in rhachidi. 
TYPE: ILLINOIS: DOUGLAS CO.: 4 miles east of Hindsboro, 
Sargent Twp. (NE!4, Sect. 3, RIOE, T14N), in open field 
at edge of road, J. E. Ebinger 6845 (EIU). This area was 
revisited on 9 October 1969 and a second collection (J. E. 
Ebinger 9282) was made at that time (EIU, ISM). 

Long, papillose-based hairs are known in a few species 
of Digitaria. The presence of these hairs is an important 
diagnostic characteristic used by Hitchcock (1935), Hen- 
rard (1950), and many others to separate the tropical Digi- 
taria horizontalis Willd. from other members of the genus. 
The hairs in this species are similar in all respects to those 

58 Rhodora [Vol. 76 

found in D. sanguinalis f. illinoensis. The two taxa are 
easily separated however, since in D. horizontalis the nearly 
glabrous spikelets are usually less than 2.4 mm long, ex- 
tremely narrow and with an acuminate apex while D. san- 
guinalis has pubescent spikelets that are longer (2.5-3.5 mm 
long) and broader and with an acute apex. The only other 
closely related taxon in which these hairs have been re- 
ported is D. adscendens (HBK) Henrard var. rhachiseta 
Henrard. The differences between this taxon and D. san- 
guinalis have been studied by Ebinger (1962, 1965). 


EBINGER, J. E. 1962. Validity of the grass species Digitaria 
adscendens. Brittonia 14: 248-253. 

. 1965. Digitaria sanguinalis in South America. 
Trans. Ill. St. Acad. Sei. 58: 255-258. 

HENRARD, J. T. 1950. Monograph of the genus Digitaria. xii + 999 
pp. Leyden, Universitare Pers Leiden. 

Hitcucock, A. S. 1935. Manual of the grasses of the United 
States. U. S. Dept. of Agric. Misc. Publ. 200. 1040 pp. (Ed. 2, 
revised by Agnes Chase, 1950). 




Growth studies on Fucus vesiculosus L., Ascophyllum 
nodosum (L.) LeJolis, Rhodymenia palmata (L.) Greville, 
Laminaria digitata (L.) Lamouroux and Laminaria sac- 
charina (L.) Lamouroux were initiated at Dover Point, 
Dover, New Hampshire and Jaffrey Point (Fort Stark), 
Newcastle Island, Portsmouth, New Hampshire in July 
1968. Dover Point is an estuarine tidal rapid area whereas 
Jaffrey Point is a semi-exposed open coastallocation. Both 
areas have been studied extensively and much is known 
of their flora and ecology (Mathieson, Hehre & Reynolds, 
in press; Mathieson, Reynolds & Hehre, in press; Reynolds, 

Approximately 20 plants of each species were tagged at 
both locations with plastic forester's tape and small white 
tags. The monthly growth of F. vesiculosus, A. nodosum 
and R. palmata was recorded by measuring their total 
length, while in L. digitata and L. saccharina growth was 
recorded by determining the monthly increase in distance 
between the intercalary meristem and paired holes (Sun- 
dene, 1964). The holes were made with a cork borer 1 inch 
above the meristem. New holes were made every 2 months, 
because the terminal portions of the lamina sloughed off 
continuously. Specimens of L. saccharina, L. digitata and 
R. palmata were transplanted from the high to the low 
current areas at Dover Point, and their growth and 
longevity was recorded. During the 16 months of the ex- 
periment, nearly all the initial plants were lost due to 
human interference, grazing of snails, wave action or 
degeneration. Consequently, it was necessary to continu- 
ously tag and measure new plants. 

The growth of Rhodymenia palmata was sporadic in the 
areas of low and high currents at Dover Point. Frag- 
mentation occurred extensively during the late winter, The 


60 Rhodora [Vol. 76 

area of low currents was not suitable for the sustained 
growth of R. palmata, for when plants were transplanted 
there they died in a few months. All specimens of R. 
palmata grew at least one inch during June 1968. 

The best growth of Fucus vesiculosus and Ascophyllum 
nodosum occurred during the summer while the poorest 
growth occurred during cold weather. The growth of F. 
vesiculosus and A. nodosum at Jaffrey Point was consistently 
lower than at Dover Point. The maximum growth of 
F. vesiculosus was 2 inches per month, and it occurred 
from May 15 to August 15, 1968, in the area of low cur- 
rents at Dover Point. 

Figures 1 and 2 show the growth of Laminaria digitata 
and L. saccharina. Because of the extreme variability in 
growth rates between individual plants, these rates are 
described only in relative terms as follows: 

(1) good growth — over 3 inches/month 
(2) average growth — 2-3 inches/month 
(3) poor growth — up to 2 inches/month. 

The growth of Laminaria digitata in the low current 
areas at Dover Point was quite variable. When trans- 
plants were made from the high to the low current areas, 
the growth was initially high, but the plants eventually 
degenerated. The probable cause is siltation and the physi- 
ological decrease of oxygen and nutrients due to the de- 
creased currents. Good growth of L. digitata occurred in 
the spring and summer at Jaffrey Point and in the area 
of high currents at Dover Point. Poor growth was particu- 
larly evident in the winter at the same locations. In general, 
L. digitata had a higher rate of growth and it was sustained 
for a longer time in the high current areas at Dover Point 
than at Jaffrey Point. 

An interesting phenomenon was observed when L. digi- 
tata was transplanted from the high to the low current 
areas at Dover Point. No dissection of the newly formed 
parts of the blades took place, and in a few months it was 
difficult to distinguish the plants from L. saccharina, ex- 
cept for the identification tags. The mechanical force of 

1974] Seaweeds — Reynolds 
H è 
5 3 
B & 3 z £ 
3 ii ii 



1- 1- | ee 
s d ; 
v 2 o 3 E $ 3 $ 
$ i d oi i $ d 
Figure 1 
RELATIVE GROWTH OF Laminaria digitata 
Poor Growth 0-2 inches per month 
Average Growth 2-3 inches per month 

Good Growth 38-54% inches per month 





high current area 
low current area 


Jaffrey tt 
Dover Pt. 


T T T "CT | 
EN . , ¢ , 
i1 : i i | i o: OB 
Figure 2 

RELATIVE GROWTH OF Laminaria saccharina 
Poor Growth 0-2 inches per month 
Average Growth 2-3 inches per month 
Good Growth 3-5 % inches per month 


1974] Seaweed — Reynolds 63 

the current was not high enough to cause the final separa- 
tion of the frond segments. 

The growth rate of L. saccharina was good for most of 
the year in the low current areas at Dover Point. A slight 
decrease in growth occurred during the winter. Good 
growth of L. saccharina occurred during the summer and 
early fall at Jaffrey Point and in the high current areas 
at Dover Point. Poor growth was evident during the late 
fall and early winter. 

Laminaria saccharina exhibited good growth at all areas 
during the summer. Laminaria saccharina dominated the 
kelp beds at Dover Point during the winter, but L. digitata 
was dominant the rest of the year. It can be concluded 
that L. saccharina is better adapted to the estuarine en- 
vironment than is L. digitata. The penetration of L. sac- 
charina into the estuary is dependent on the presence of 
currents to compensate for the reduced salinities. 


MATHIESON, A. C., E. HEHRE & N. B. REYNOLDS. Investigations of 
New England marine algae I. A floristic and descriptive eco- 
logical study of the marine algae of Jaffrey Point, New Hamp- 
shire. Nova Hedwigia, In press. 

MATHIESON, A. C., N. B. REYNOLDS & E. HEHRE. Investigations of 
New England marine algae. II. Distribution of benthonic marine 
algae in the Great Bay Estuary System. Nova Hedwigia, In 

REYNOLDS, N. B. 1971. The ecology of a New Hampshire estuarine 
tidal rapid. Ph.D. Thesis, University of New Hampshire Li- 
brary, 101 pp. 

SUNDENE, O. 1964. The ecology of Laminaria digitata in Norway 
in view of transplant experiments. Nytt Mag. Bot. 11: 83-107. 





Heretofore, knowledge of the cytology of Kosteletzkya 
Presl has consisted of a single chromosome count from 
K. hastata Presl (Skovsted, 1941). This note corrects the 
number erroneously reported by Skovsted and adds counts 
from two African species, K. adoensis (Hochst. ex A. Rich.) 
Mast., and K. buettneri Gürke, and four American species, 
K. coulteri A. Gray, K. pentasperma (Bertero ex DC.) 
Griseb. K. paniculata Benth., and K. virginica (L.) Presl 
ex A. Gray (Table 1). 

Counts were made from pollen mother cells of buds col- 
lected in the wild or taken from plants grown from seeds 
collected in the wild. Methods of fixation, preparation, and 
documentation are those reported by Bates and Blanchard 
(1970). Voucher specimens have been deposited in the 
L. H. Bailey Hortorium (BH). 

The species counted represent a rather wide range of 
morphological diversity in the genus as well as a con- 
siderable geographical range. Kosteletzkya paniculata be- 
longs to the specialized sect. Orthopetalum Benth. of 
western Mexico, which is notable in its convolute, tubular 
corolla and its exserted staminal column. The remaining 
species, both African and American, belong to the wide- 
spread sect. Kosteletzkya, but within that section repre- 
sent a diversity of forms. The fact that counts made from 
all these species are the same, n = 19, suggests that the 
number in the genus may be constant, At variance with 
this conclusion is Skovsted's report of 2n = 34 in K. has- 
tata, a species closely related to K. pentasperma. How- 
ever, an examination of the illustration by Skovsted (1941, 
fig. 99) shows 38 chromosomes rather than the reported 
34. It may therefore be assumed that the gametic chromo- 
some number of K. hastata is also » — 19. 


1974] Kosteletzkya — Blanchard 65 

The error that entered Skovsted's paper is of further 
interest because he used the erroneous count to support 
Hochreutiner's suggestion (1900) that Kosteletzkya could 
be related to Hibiscus sect. Pterocarpus Garcke ex 
Hochreutiner, a section known cytologically only from 
H. vitifolius L. with a chromosome number of n = 17 
(Skovsted, 1935, 1941). There are ample reasons on 
morphological grounds for relating Kosteletzkya and 
Hibiscus sect. Pterocarpus, e.g., they share depressed, 
pentagonal, 5-crested capsules, the valves of which fall 
away completely from the floral axis at maturity, as well 
as similarities in calyx, involucre, and vegetative char- 
acters. Now, however, any proposal to unite the two taxa 
must take into account the difference in chromosome 


Field collections were aided by funds from National 
Science Foundation Grant GB-8759 to David M. Bates. 
I also wish to thank Paul A. Fryxell for providing trans- 
portation in Mexico. 


Bates, D. M., & O. J. BLANCHARD, JR. 1970. Chromosome numbers 
in the Malvales. II. New or otherwise noteworthy counts rele- 
vant to classification in the Malvaceae, tribe Malveae. Amer. 
J. Bot. 57: 927-934. 

HOCHREUTINER, B. P. G. 1900. Revision du genre Hibiscus, An- 
nuaire Conserv. Jard. Bot. Genéve 4: 23-191. 

SKOVSTED, A. 1935. Chromosome numbers in the Malvaceae. I. 
J. Genet. 31: 263-296. 

1941. Chromosome numbers in the Malvaceae. II. 

C. R. Lab. Carlsb. Physiol. 23: 195-242. 

66 Rhodora [Vol. 76 

Table 1. Chromosome counts in Kosteletzkya 

Species n-number Locality and Collector 
K. adoensis 19 Cultivated, BH-69:555. Seed 
source — Angola: Huambo 

Distr., Instituto de Investigação 
Agronómica. de Angola 

K. buettneri 19 Cultivated, BH-72:100. Seed 
source — Zambia: “C Pro- 
vince," Robinson 6706 
K. coulteri 19 Mexico: Sinaloa, Fryxell, Bates 
& Blanchard 1544 
K. pentasperma 19 Mexico: Jalisco, Fryxell, Bates 
& Blanchard 1610 
19 Mexico: Michoacan, Fryzell, 

Bates & Blanchard 1650 
19 Mexico: Nayarit, Fryxell, Bates 
& Blanchard 1563 

K. paniculata 19 Cultivated, BH-71:119. Seed 
source — Mexico: Jalisco, 
Fryxell, Bates & Blanchard 

K. virginica 19 USA: Florida, Collier Co., 

Blanchard & Blanchard 303 
19 USA: New Jersey, Ocean Co., 
Blanchard & Blanchard 284 
19 Cultivated, BH-69:430. Seed 
source — USA: Texas, Jeffer- 
son Co., Knight 88 




The author has been working in collaboration with 
Richard A. Howard of the Arnold Arboretum and George 
R. Proctor of the Institute of Jamaica towards a revision 
of Britton and Millspaugh’s “Bahama Flora," published 
first in 1920 and reprinted without changes in 1962. A 
recent paper by the three of us (Gillis, Howard, and 
Proctor, 1973) has reported additions to the flora of the 
Bahamas based on personal collections, new collections 
available to us, and reports from the literature. 

The present paper offers updated annotations on the 
correct scientific names to be applied to the species listed 
by Britton and Millspaugh whose work was completed 
under the provisions of the American Code of Botanical 
Nomenclature. We encourage others to use this list and its 
supporting bibliography, and to report additional changes 
or additions to me. We would particularly value contribu- 
tions which specialists in non-vascular plants could make 
to the listings in Britton and Millspaugh for mosses and 
liverworts, algae, fungi, slime molds, and lichens. Our 
efforts will consider only the flowering plants and pteri- 
dophytes. We therefore encourage workers in these other 
fields of botany to update knowledge of their disciplines 
in the Bahamas too. 

The value of lists of nomenclatural corrections has 
been evident to us in the work of Alain (1965) on the flora 
of Puerto Rico and the Virgin Islands, Additional nomen- 
clatural changes have been obtained from monographs or 
the recent floras of Jamaica (Adams, 1971) and Barbados 
(Gooding et al., 1965). In all of these, however, the rea- 
sons for the changes of names are not always explained, 
requiring the curious or careful worker to reinvestigate 
each and every problem. I believe the explanations which 
aecompany the changes reported here will be of value to 


68 Rhodora [Vol. 76 

other workers in subtropical New World floras and par- 
ticularly in the West Indies. Reasons given for changes are 
usually: use of an earlier epithet, avoidance of a tautonym, 
replacement of a later homonym, correct application of a 
name, etc. When name changes have been made by others 
than myself, I have cited the publication upon which this 
information was based. Conversely, when I have had 
reason not to accept a particular treatment, then reasons 
have been stated. 

The listing is by no means complete. Certain groups 
within the Boraginaceae, Rubiaceae, Loranthaceae, and 
Asclepiadaceae need intensive study and many more obser- 
vations in the field, especially in such genera as Helio- 
tropium, Borreria, Dendropemon, Phthirusa, and Cynan- 
chum. A new look must be given to the genus Agave for 
the West Indies, inasmuch as the most recent treatment 
is now 60 years old (Trelease, 1913). Other groups in- 
cluding the ferns are being studied by others at the present 
time and we await publication of the conclusions of these 

The Turks and Caicos Islands are politically a separate 
Crown Colony from the newly independent Commonwealth 
of the Bahama Islands. Britton and Millspaugh included 
all within the Bahama Flora and such is the geographical 
sense employed here in that it includes the Turks and 
Caicos Islands as well. 

For ease of reference this paper will follow the order of 
species presented in Britton and Millspaugh’s Bahama 
Flora, which will be Monocotyledons first, followed by the 
Dicotyledons, and then the Gymnosperms. Again for 
brevity, Britton and Millspaugh’s flora is referred to by 
the expression B&M in the text of this paper. The figure 
in the left margin refers to the page in B&M on which the 
taxon in question is discussed. The name following the 
arrow (—) is the name considered to be correct. These 
names are not necessarily nomenclatural or taxonomic 
equivalents, hence the reason for not employing an equals 

1974] Bahama Flora — Gillis 69 

sign (—). In some instances, B&M misidentified the plant 
or used a binomial incorrectly. Where monographs or 
significant studies support the use of names as they were 
employed in B&M, such references are cited under the 
generic name, e.g., Guaiacum (Porter, 1972). 

In a few instances where the name changes have involved 
closely allied genera, or where the existing keys in B&M 
are wholly inadequate, particularly for use in the field, I 
have provided new keys. They are designed to employ 
characters of the floral, fruiting, and vegetative portions 
of the plant so that they can be of optimum use in working 
with specimens, whether living or in the herbarium, and 
whether reproductive or vegetative. 


I should like to acknowledge the help of my associates Dr. 
Howard and Mr. Proctor in drawing up this list, of Dr. 
Carroll E. Wood, Jr. and Mr. George N. Avery, who read 
the manuscript and made valuable suggestions, of my col- 
league Dr. Kenneth R. Robertson who gave advice, and of 
the many workers who have studied genera that have 
representatives in the Bahamas. Special appreciation is 
due to the librarians at the Arnold Arboretum and Gray 
Herbarium of Harvard University and for the use of the 
bibliographic files of the Generic Flora of the Southeastern 
United States project, made available by Dr. Wood. I should 
also like to acknowledge with thanks the continuing co- 
operation of the personnel of the Ministry of Agriculture 
and Fisheries, Commonwealth of the Bahama Islands. 

This study was made while the author was a Research 
Fellow at the Arnold Arboretum and I express my grati- 
tude for this opportunity. A generous gift from an anony- 
mous donor interested in the work on the Bahama flora. 
made this study possible. 




Rhodora [Vol. 76 

Typha angustifolia > Typha domingensis (Pers.) 
Kunth, These are two different valid species accord- 
ing to Hotchkiss and Dozier (1949) with only Typha 
domingensis occurring in the Bahamas. 


Potamogeton heterophyllus > Potamogeton gramineus 
L. var. gramineus. On the basis of both herbarium 
specimens and abundant fresh material from a pond 
on San Salvador Island (Gillis 8864), I have con- 
sidered all the Potamogeton material on the Islands 
to be the same (see Ogden, 1948). 

The question of good characters, reliable under all 
conditions, for separating Ruppia maritima from 
R. cirrhosa, is still open. Various treatments have 
been consulted (Fernald and Wiegand, 1914; Hag- 
strom, 1916; McCann, 1945; Setchell, 1946; Reese, 
1962a and b; Gamerro, 1968; and Schwanitz, 1967) 
for specimens collected in the Bahamas. For the 
moment, it appears that both species seem to be 
present. Therefore, R. cirrhosa is an addition to the 
flora, rather than merely a different name for the 
species in B&M. Collectors are urged to look for R. 
cirrhosa inasmuch as flowering and fruiting material 
are as yet unknown in the Bahamas. An adaptation 
of Gamerro's key follows: 

Peduncle of inflorescence more than 10 cm. (8-60 
em.), coiled until anthesis; anther sacs reniform, 
1.7 mm. long; fruits rounded at apex; leaves 1 mm. 
broad; leaf epidermal cells 16-194; 2n = 40 .... 
TEN Ruppia cirrhosa (Petagna) Grande. 
Peduncle of inflorescence less than 5 cm., not coiled 
in spiral until anthesis; anther sacs elliptic, 0.6-0.7 
mm. long; fruits irregularly pointed; leaves 0.5 





Bahama Flora — Gillis 71 

mm. broad; epidermal cells of leaves 12.5-16.0,5 
Oe ee et Mert a tad Ruppia maritima L. 


Cymodocea manatorum —> Syringodium filiforme Kütz. 
This change of name follows Dandy and Tandy (1939) 
and den Hartog (1970). 
Halodule wrightii Aschers. is indeed in the Bahamas, 
but so is H. beaudettei (den Hartog) den Hartog 
(1964). These two species can be separated as fol- 

Leaf tips bicuspidate; leaf blades 1/3-4/5 mm. 
wide; leaves with linear teeth, concave on the inner 

side ... Halodule wrightii Aschers. 
Leaf tips tricuspidate; leaf blades 2/3-114 mm. 
wide; leaves with linear lateral teeth ... Halodule 

beaudettei (den Hartog) den Hartog. 

Halophila — consult Hartog (1959). 


Echinodorus cordifolius — Echinodorus berteroi 
(Spreng.) Fass. (See Fassett, 1955). 


There is some question among agrostologists whether 
Schizachyrium should be segregated from Andropogon 
as it was in B&M. Chase (1951) joins the two genera. 
For the moment, however, I shall follow treatments 
by Gould in maintaining them as separate genera 
(see Gould, 1967, 1968, and 1969). 

Nazia aliena > Tragus berteronianus Schult. Tragus 
is conserved over Nazia. The species epithet changed 
from aliena to berteronianus because the type of 
Lappago alienus Spreng., the basionym, is referred 
to Pseudechinolaena. polystachya (H.B.K.) Stapf, and 
not to this Bahamian plant (Chase, 1951). 











Rhodora [Vol. 76 

Syntherisma filiformis > Digitaria panicea  (Sw.) 
Urb. The species on which the epithet filiformis is 
based is a different species from the one of concern 
here. The earliest name for this species is Milium 
panicea Sw. which was transferred to Digitaria by 
Urban. The combination was made twice in the 
same year (1920), by Urban in February and by 
Fernald in June, The Urban combination obviously 
has priority. Digitaria is an earlier generic name 
than Syntherisma. 

Syntherisma digitata — Digitaria horizontalis Willd. 
Milium digitatum Sw. (1788) is the earliest basionym, 
but is is preoccupied in Digitaria by D. digitata Buse 
(1854). The next earliest name available is D. kori- 
zontalis Willd. (1809). 

Syntherisma sanguinalis > Digitaria sanguinalis (L.) 

Valota insularis > Trichachne insularis (L.) Nees. 
Valota is inadequately published (Chase, 1951). 
Paspalum poiretii R. et S. ^ Paspalum sagetii Chase. 
Paspalum caespitosum — Paspalum poiretii R. et S. 
The work of Chase (1929) sorts out the names of 
these Paspalum species by typifying them. It is thus 
apparent that, although the name P. poiretii is still 
used in the Bahama flora, it must be applied to a 
different species from the one to which it was as- 
signed by B&M. 

Paspalum  portoricense ^ Paspalum molle Poir. in 
Lam. The change is required because of an earlier 
epithet (Chase, 1951). 

Paspalum simpsonii > Paspalum blodgettii Chapm. 
Paspalum blodgetti is an earlier name for P. simp- 
sonii Nash, and not a synonym of P. caespitosum 
Chase, 1951). 

Paspalum glabrum — Paspalum laxum Lam. The 
change is required because of an earlier epithet 
(Chase, 1951). 

Paspalum distichum L. is attributed by B&M to the 









Bahama Flora — Gillis T8 

Pugil. Pl. Jam., one of the dissertations of Linnaeus 
(1759). It actually appeared earlier in the same 
year in the Systema Naturae, ed 10, and should be 
so attributed. 

Panicum barbinode > Panicum purpurascens Raddi. 
The change is required because of an earlier epithet 
(Hitchcock, 1936). 

Panicum distantiflorum — Setaria distantiflorum (A. 
Rich. in Sagra) Pilger. 

Panicum chapmani — Setaria chapmanii (Vasey) 
Pilger. Rominger (1962) has shown that these two 
species properly belong in Setaria. 

Panicum dichotomiflorum — Panicum _ bartowense 
Scribn. et Merr. These names were treated as synony- 
mous by B&M. Hitchcock (1935) and Chase (1951) 
felt that they are separate species and that the Baha- 
mian plant is P. bartowense, Fernald (1934) treated 
the taxon as P. dichotomiflorum var. bartowense 
Schribn. et Merr.) Fern. 

Panicum coerulescens > Panicum coerulescens Hack. 
ex Hitehe. This change is simply a corrected spelling. 
Oplismenus hirtellus (L.) R. & S. 1817 — Oplismenus 
hirtellus (L.) Beauv., Ess. Agrost. 54, 168. 1812. An 
author change is needed for this earlier publication 
of the binomial. 

Chaetochloa geniculata — Setaria geniculata (Lam.) 

Chaetochloa setosa — Setaria setosa (Sw.) Beauv. 
Chaetochloa macrosperma — Setaria macrosperma 
(Scribn. et Merr.) K. Schum. Setaria is the older 
and preferred name over Chaetochloa. Setaria Beauv. 
is conserved over Setaria Michx. 

Cenchropsis mysuroides — Cenchrus mysuroides 
H.B.K. Cenchropsis is not now considered distinct 
from Cenchrus (Chase, 1936 and Delisle, 1963). 
Cenchrus carolinianus ^ Cenchrus incertus M. A. 
Curtis. The use of Cenchrus carolinianus in B&M 
was a misapplication of the name. That name is 








Rhodora [Vol. 76 

now considered a synonym of C. longispinus, but is 
correctly used for a plant not found in the Bahamas. 
The plant in question is C. incertus which is wide- 
spread in the Bahamas (DeLisle, 1963). 

Cenchrus microcephalus > Cenchrus incertus M. A. 
Curtis. DeLisle (ibid.) felt that there was so little 
difference between the *endemic" race of Cenchrus 
with small fruits and the more widespread species 
that he treated them as conspecific. The small fruited 
form should be considered an insular form. 
Cenchrus viridis —^ Cenchrus brownii R. et S. This 
name change is required because of an earlier epithet 
(DeLisle, ibid.). 

Stenotaphrum — Consult Sauer, 1972. 

Aristida gyrans > Aristida vilfifolia Henr. After 
publication of B&M, Aristida vilfifolia was segregated 
as a distinct species from A. gyrans which was the 
name in B&M (Henrard, 1926-33; 1929-33). It is 
the segregate which indeed is the one in the Bahama 

Aristida scabra > Aristida ternipes Cav. This change 
is required because of an earlier epithet. 

Sporobolus indicus — Sporobolus jacquemontii Kunth. 
Sporobolus berteroanus — Sporobolus indicus (L.) R. 
Br. In the Sporobolus indicus complex, there are two 
Caribbean species which are often confused: S. jac- 
quemontii, which has an obtuse upper glume and is 
a fully tropical species, and S. indicus, which has an 
acute upper glume and is a more subtropical species. 
According to the description in B&M, these taxa have 
been confused (Clayton, 1964 and personal communi- 

Sporobolus atrovirens — Sporobolus bahamensis Hack. 
Sporobolus atrovirens is a different plant from the one 
in the Bahamas; it is a Mexican plant, is a perennial, 
and has smaller spikelets and glumes than S. baha- 
mensis. The latter is a Bahamian endemic and an 








Bahama Flora — Gillis 75 

Sporobolus argutus > Sporobolus pyramidatus (Lam.) 
Hitehe. This name change is required because of an 
earlier basionym. The citation for Domingan drop- 
seed-grass should be Sporobolus domingensis (Trin.) 
Kunth, Rév. Gram. 1, Suppl.; xvii. 1830, a citation 
three years earlier than that cited in B&M. 
Capriola dactylon > Cynodon dactylon (L.) Pers. 
Cynodon is conserved over Capriola, 

Chloris polydactyla > Chloris dandyana Adams. 
Adams (1971) published a new name for this plant 
without giving the background reasons for the illegiti- 
macy of the original name. The background is as 
follows: There is a Jamaican plant with the legiti- 
mate name Andropogon barbatus L. Some time after 
describing this Jamaican grass, Linnaeus applied the 
same binomial to an East Indian grass. The epithet 
barbatus (referring to the East Indian species) was 
transferred to Chloris by Swartz, an act which pre- 
vents the transfer of the Jamaican species epithet to 
Chloris. In the meantime, Linnaeus proposed a super- 
fluous name for the first Andropogon barbatus, i.e., 
A. polydactylos. Swartz’s transfer of this super- 
fluous name to Chloris is illegitimate; hence Adams’s 
publication of a new name for the species in the West 
Indies. (I should like to acknowledge personal com- 
munication from C. D. Adams and J. E. Dandy for 
the historical background on this matter.) 
Phragmites phragmites ^5 Phragmites australis (Cav.) 
Trin. ex Steud. Clayton (1968) pointed out that the 
correct name for the pan-tropical reed is the com- 
bination used above. 

Diplachne fascicularis > Leptochloa fascicularis (Lam.) 
A. Gray. Current views of agrostologists suggest that 
Diplachne should be contained within Leptochloa. 
Eragrostis cilaris (L.) Link 1827  Eragrostis cili- 
aris (L.) R.Br. in Tuckey, 1818. Chase (1951) re- 
corded the earlier publication of this combination. 








Rhodora [Vol. 76 - 

Uniola virgata > Leptochloópsis virgata — (Poir.) 
Yates. Yates (1966) has subdivided Uniola into 
Uniola, Chasmanthium, and Leptochlodpsis on the 
basis of a number of anatomical, cytological, morpho- 
logical, and embryological characters. Because this 
species of spike-grass is considered close to Lepto- 
chloa, Yates erected Leptochlodpsis to contain it and 
another species from Ecuador. 


Cyperus paniculatus > Cyperus polydactylos Rottb. 
These names originate from the same date and are 
considered to be synonymous. Corcoran (1941) chose 
C. polydactylos to be the inclusive name. The Bahama 
populations might be designated as C. polydactylos 
var. texensis (Torr.) Fern. (McLaughlin, 1944). 

Cyperus cuspidatus ^ Cyperus aristatus Rottb. True 
C. cuspidatus does not occur in the Bahamas, but in 
Cuba. (See Horvat, 1941; McLaughlin, 1944). 
Cyperus pseudovegetus > Cyperus distinctus Steud. 
Cyperus pseudovegetus was a name misapplied to the 
plant of the Bahamas (O’Neill, 1939). 

Cyperus brunneus — Cyperus planifolius L. C. Rich. 
This name change is required by an earlier epithet 
(McGivney, 1938; McLaughlin, 1944). 

Cyperus ferax — Cyperus odoratus L. var. odoratus. 
O'Neill (1940) demonstrated that these two names 
apply to the same species, and that C. odoratus is the 
earlier name. He further thought that the typical 
variety was the one found in the Bahamas. 

According to the treatment by McLaughlin (1944), 
the populations of Cyperus filiformis Sw. in the 
Bahamas are of the typical variety (var. filiformis). 
Stenophyllus wilsonii > Bulbostylis floccosa (Griseb.) 
Clark in Urb. Stenophyllus is merged with Bulbo- 
stylis in the treatment by Kral (1971). Bulbostylis 
floccosa is the earliest name. It is not endemic to the 











Bahama Flora — Gillis T4 

Bahamas as stated in B&M, but also is found in Cuba 
and Hispaniola. 

Fimbristylis diphylla — Fimbristylis dichotoma (L.) 
Vahl. This change is required because of an earlier 

According to B&M, the record of Fimbristylis hirta 
in the Bahamas is very tenuous. Neither Kral nor 
I have seen any material from the Bahamas. If in- 
deed it does occur there, its more appropriate name 
should be F. squarrosa Vahl (see Kral, 1971). 
Abildgaardia monostachya > Abildgaardia ovata 
(Burm. f.) Kral. This change is required because of 
an earlier epithet (Kral, ibid.). 

There are cyperologists who wish to merge Dichro- 
mena with Rhynchospora. On the basis of the striking 
white bracts and insect pollination characteristic of 
Dichromena, I am, for the moment at least, retaining 
it as distinct. 

Rhynchospora cyperoides > R. cyperoides (Sw.) 
Mart. var. cyperoides. 

Rhynchospora tracyi —^ Rhynchospora cyperoides var. 
triceps (Vahl) Bock. This treatment follows that of 
Kükenthal (1949). 

Rhynchospora perplexa > Rhynchospora microcarpa 
Baldw. ex Gray. Gale (1944)maintained that true 
R. perplexa was not in the Bahamas and that all the 
material which Britton and Millspaugh called by this 
name was misdetermined. 

Rhynchospora bahamensis Britt. ~ Rhynchospora lin- 
deniana var. bahamensis (Britt.) Gale. Gale's treat- 
ment (ibid.) of the West Indian Rhynchospora species 
reduced Britton’s species to a variety of the more 
widespread R. lindeniana. 

Rhynchospora marisculus > Rhynchospora odorata 
C. Wright ex Griseb. Gale (ibid.), in examining the 
same material which Britton and Millspaugh did, 
determined that the use of R. marisculus in B&M was 
a misapplication of the name. 






Rhodora [Vol. 76 

Mariscus jamaicensis > Cladium jamaicense Crantz. 
Most treatments now segregate Cladium from 

Scleria. Consult Jackson, 1949. 

Thrinax parviflora > Thrinax floridana Sarg. Thri- 
nax parviflora is an endemic species of Jamaica. 
Furthurmore, the treatment of Thrinax by Read in 
Adams (1971) will be superseded now that additional 
type material has been examined. The correct name 
for the small thatch of South Florida and the northern 
West Indies with leaf segments broadest at the point 
of fusion, with scattered, fimbriate, centrally trans- 
lucent seales, and glabrous axes should be T. floridana 
(Read, personal communication). 
Thrinax parviflora > Thrinax floridana Sarg. Thri- 
(Jacq.) L. H. Bailey. True C. argentea may exist in 
the southern Bahamas, but this has yet to be demon- 
strated. It is now treated as being indigenous to 
Hispaniola. (See Moore, 1963). 
Paurotis wrightii > Acoelorrhaphe wrightii (Griseb. 
ex H. Wendl.) H. Wendl. ex Becc. The preferred 
name for the Paurotis palm, Spanish-top, or Ever- 
glades palm is now Acoelorrhaphe (Moore, 1963). 
Pseudophoenix vinifera > Pseudophoenix  sargentii 
subsp. saonae (Cook) Read. True P. vinifera is in- 
digenous to Hispaniola and is not known from the 
Bahamas. (Read, 1968). 

Commelina longicaulis ^ Commelina diffusa Burm. f. 
Commelina elegans —^ Commelina virginica L. Name 
changes are made here in accord with the treatment 
by Brashier (1969). The following key may be more 
easily used than that in B&M: 

Plant annual, trailing; spathes not united at the 
base, margins ciliate; internodes glabrous; sta- 












Bahama Flora — Gillis 79 

minodes Qoi m Commelina diffusa Burm. f. 
Plant perennial, tufted and erect; spathes united at 
the base, margins membranous; internodes puberu- 
lent; staminodes 3 ...... Commelina virginica L. 

Rhoeo discolor ^ Rhoeo spathacea (Sw.) Stearn. This 
name change follows Stearn (1957) who discovered 
that the earlier epithet of Swartz applies to this 


Cordyline guineensis 5 Sansevieria hyacinthoides 
(L.) Druce. This change follows Wijnands (1973). 

Smilax. Consult Coker, 1944. 


Atamosco rosea — Zephranthes rosea Lindl. 
Atamosco cardinalis > Zephyranthes cardinalis C. 
Wright. Zephyranthes is conserved over Atamosco. 
Hymenocallis caymanensis ^ Hymenocallis latifolia 
(Mill.) Roemer. This change follows the treatment 
by Sealy (1954). 


Carteria corallicola — Basiphyllaea corallicola (Small) 
Ames. Carteria is a later homonym for a genus of 
algae. Basiphyllaea was erected as a genus to correct 
this situation. 

Vanilla eggersii — Vanilla dilloniana Correll. 
Vanilla articulata — Vanilla barbellata Reichb. f. This 
treatment of Vanilla in the Bahamas follows Cor- 
rell (1950). 

Pelexia adnata — Spiranthes adnata (Sw.) Benth. 
Although Pelexia is a conserved name, it is also a 
generic synonym of Spiranthes (Garay and Sweet, 

Pelexia setacea > Eltroplectris calcarata (Sw.) Garay 











Rhodora [Vol. 76 

et Sweet. Luer (1972) changed Pelexia setacea to 
Centrogenium setaceum but Garay and Sweet (1972) 
maintained that this combination is based on an ille- 
gitimate name and therefore should be Eltroplectris 
calcarata (Sw.) Garay et Sweet. 

Ibidium tortile > Spiranthes tortilis (Sw.) L. C. 
Rich. Jbidiwm is considered to be a generic synonym 
of Spiranthes. (Garay & Sweet, 1972). 

Ibidium lucayanum — Spiranthes polyantha Reichb. f. 
These names are considered synonymous by Luer 

Stenorrhynchus lanceolatus — Spiranthes lanceolata 
(Aubl) Léon. Stenorrhynchus is now treated as a 
generic synonym of Spiranthes. (See Dunsterville 
and Garay, 1965). 

Ponthieva brittonae > Ponthieva racemosa var. brit- 
tonae (Ames) Luer. Two varieties of Ponthieva 
racemosa occur in the Bahamas. One is the typical 
variety (var. racemosa) as treated by Luer (1972) 
and reported by him as new to the flora. Var. brit- 
tonae is the reduction made by Luer of the name in 

Physurus querceticola —^ Erythodes querceticola 
(Lindl.) Ames. Physurus is considered to be a gene- 
ric synonym of Erythodes (Garay and Sweet, 1972). 
Polystachya minuta — Polystachya flavescens (Lindl.) 
J. J. Smith. Although the epithet minuta (from 
Epidendrum minutum Aubl.) is the oldest name 
available, it is preoccupied in Polystachya, and hence, 
P. flavescens must be used. 

Spathiger | rigidus > Epidendrum rigidum Jacq. 
Spathiger is considered a generic synonym of Epi- 

Auliza nocturna > Epidendrum nocturnum Jacq. 
Auliza is considered a generic synonymn of Epiden- 
drum (Dunsterville and Garay, 1965; Luer, 1972). 

There is much discussion among orchid taxonomists 
whether Encyclia should be segregated from Epiden- 










Bahama Flora — Gillis 81 

drum or not. For the moment, I accept the treat- 
ment of Dressler (1961; 1966) and Beckner (1970) 
in maintaining Encyclia at the generic level as in 

Encyclia rufa > Encyclia bahamensis (Griseb.) Britt. 
et Millsp. Specimens labeled E. rufa from the Ba- 
hamas are now all best considered E. bahamensis. 
Encyclia rufa is, as B&M suspected, still not known 
from the archipelago. 

Encyclia diurna — Encyclia hodgeana (A. D. Hawkes) 
Beckner. This name change is required because of 
more complete understanding of the taxa involved. 
The species previously called E. diurna or E. gracilis 
is, in fact, a hybrid between E. hodgeana and E. 
tampensis. (See Beckner, 1970). 

Epicladium boothianum — Encyclia boothiana var. 
erythronioides (Small) Luer. Epicladiwm is a generic 
synonym of Encyclia. 

Anacheilium cochleatum — Encyclia cochleata var. 
triandra (Ames) Dressler. Anacheiliwm is a generic 
synonym of Encyclia. 

Laeliopsis domingensis ^ Broughtonia lindenii (Lindl.) 
Dressler. This change is made in accord with the 
treatment by Dressler (1961). 

Limodorum simpsoni — Calopogon tuberosus (L.) 
Britten, Stearns, and Poggenberg. This treatment 
follows Luer, 1972. 

Picrodendron macrocarpum — Picrodendron baccatum 
(L.) Krug et Urb. in Engl. I presently have Picro- 
dendron under study. It appears that this genus 
should be placed within the Euphorbiaceae. There 
is likely only one species, which would be the typical 
one, P. baccatum. 

Trema lamarckiana — Trema lamarckianum (R. et S.) 
Blume. The gender of Trema should be neuter. 


Rhodora [Vol. 76 


105 Ficus jacquinifolia > Ficus perforata L. 

Fieus brevifolia — Ficus citrifolia Mill. Recent treat- 
ments of Ficus (DeWolf, 1969 and Condit, 1969) have 
equated F. perforata with F. jacquinifolia A. Rich. 
in Sagra. Moreover, they have also equated F. brevi- 
folia and F. citrifolia. B&M’s F. brevifolia Nutt. 
(lectotype at BM) must become F. citrifolia Mill., the 
earlier name (lectotype at BM, chosen by Dandy). I 
have further studied F. perforata which DeWolf and 
Condit have given as the correct name for F. jac- 
quinifolia (which itself should be written jacquinii- 
folia). The Linnaean name must be typified by an 
illustration in Plumier associated with the disserta- 
tion on the plants of Surinam (1775). Although there 
is a specimen of F. perforata at LINN (Savage Cata- 
log No. 1240: 9), the only reference in the Disserta- 
tion (and the later Amoenitates) in this context is 
that to Plumier. The specimen at LINN appears to 
be of the tree of Central America and northern South 
America, F. pertusa L. f. I should like to acknowledge 
the consultation and discussion with Dr. Gordon De- 
Wolf which proved very valuable in application of 
names in Ficus. A key to the Bahama material fol- 

1. Syconia sessile .......... Ficus aurea Nutt. 
1. Sycania stalked 
2. Syconia 7-10 mm. in diameter; petioles less 
than 1 cm.; blades obovate; lateral veins 

flush with lower surface; leaf base cuneate 
Lee eee eee eee ee eee Ficus perforata L. 

2. Syconia 8-18 mm. in diameter; petioles 
longer than 1 cm.; blades ovate; lateral veins 

+ prominent, somewhat elevated on lower 
surface; leaf blade base rounded ......... 
Lee eee ee eee eee eee Ficus citrifolia Mill. 









Bahama Flora — Gillis 83 


Generic study of Dendropemon, Phthirusa, and Phora- 
dendron is needed before good species concepts and 
correct nomenclature emerge. 

Aristolochia passifloraefolia — Aristolochia passiflori- 
folia A. Rich. in Sagra. With the exception of the 
orthograhic change (above) made in accord with 
rules adopted at the Seattle Congress, the names for 
Aristolochia species in the Bahamas remain as they 
were in B&M (Pfeifer, 1966; 1970). 

Persicaria hydropiperoides > Polygonum hydropipe- 
roides Michx. 
Persicaria punctata — Polygonum punctatum Ell. 
Persicaria portoricensis Polygonum densiflorum 
Meissn. Persicaria is treated by Graham and Wood 
(1955) as a section of Polygonum. 
Fagopyrum fagopyrum —> Fagopyrum esculentum 
Moench. This change is necessary in order to avoid a 
Coccolobis laurifolia — Coccoloba diversifolia Jacq. 
Coccolobis bahamensis — Coccoloba tenuifolia L. 
Coccolobis diversifolia —^ Coccoloba swartzii Meisn. 
Coccoloba is conserved over Coccolobis. All Coccoloba 
species therefore should change spelling to conform. 
The 3 species mentioned above are changed further 
in accord with Howard (1957), From Howard's 
treatment also comes the change of citation for 
Coccoloba uvifera (L.) L., Syst. Nat., ed. 10, p. 1007. 

Dondia linearis > Suaeda linearis (Ell.) Moq. 
Dondia fruticosa —> Suaeda fruticosa (L.) Forsk. 
Dondia insularis ^ Suaeda insularis (Britt.) Urb. et 
Ekm. Dondia is considered to be a taxonomic synonym 












Rhodora [Vol. 76 

of Suaeda, Suaeda Forsk. ex Scopoli remains in the 
list of conserved names even though the reason for 
its inclusion is no longer valid. 


Amaranthus gracilis > Amaranthus viridis L. Reed 
(1968) and Merrill (1936) considered these species 
to be taxonomic synonyms. Amaranthus viridis L. 
is the earlier name. 

Centrostachys indica — Achyranthes aspera var. in- 
dica L. Centrostachys seems better treated as a 
generic synonym Achyranthes. 

Achyranthes maritima — Alternanthera maritima St. 

Achyranthes polygonoides — Alternanthera  polygo- 
noides (L.) R. Br. 

Achyranthes repens —> Alternanthera pungens H.B.K. 
This treatment follows Reed, 1968. The epithet 
pungens is used for the last species rather than the 
earlier epithet repens because Achyranthes repens is 
a nomen confusum according to Reed. 

Philoxerus remains the same except for citing an 
earlier date of publication: Philoxerus vermicularis 
(L.) R. Br. Prodr. Fl. Nov. Holl. 416. 1810. 

Iresine celosia — Iresine diffusa Humb. et Bonpl. ex 
Willd. (Sp. Pl., ed. 4, 4: 765. 1806). Iresine celosia 
is an illegitimate name. 

Boerhaavea coccinea Mill. > Boerhavia coccinea Mill. 
Boerhaavea erecta L.— Boerhavia erecta L. Boer- 
havia is the original spelling. 

Torrubia obtusata > Guapira obtusata (Jacq.) Little. 
Torrubia cokeri > Guapira obtusata (Jacq.) Little. 
Torrubia longifolia > Guapira longifolia (Heimerl) 

Torrubia bracei > Guapira bracei (Britt.) Little. If 
one considers that blollies are similar to Pisonia 









Bahama Flora — Gillis 85 

species, then Torrubia merges with Pisonia. If one 
wishes to recognize both as distinct genera, then one 
must take up Guapira, an earlier name than Torrubia. 
The Committee for Spermatophytes rejected a pro- 
posal to conserve Torrubia, hence a paper by Little 
(1968) which made the appropriate transfers to 
Guapira, It is currently felt that Torrubia cokeri is 
synonymous with Guapira obtusata; it has been so 
treated here. This is an exceedingly variable species 
with different forms having been named in the past. 

Portulaca phaeosperma —> Portulaca rubricaulis H.B.K. 
Legrand (1952) chose the earlier name for this 

Castalia pulchella > Nymphaea pulchella DC. Wood 
(1959) has treated Castalia as a subgenus of Nym- 

Ocotea coriacea > Nectandra coriacea (Sw.) Griseb. 
Recent treatments have merged Ocotea and Nectandra 
(Wood, 1958; Allen, 1966). 
Persea pubescens Persea palustris (Raf.) Sarg. 
Kopp (1966) pointed out that palustris is an earlier 
epithet than pubescens for this species. 

Cassytha americana —> Cassytha filiformis L. This 
change is necessitated because filiformis is the earlier 

Hitchcock (1945) recognized several varieties of 
Lepidium virginicum. If this treatment is followed, 
the Bahama one is var. virginicum. 
Sinapis arvensis Brassica kaber (DC.) L. C. 
Wheeler. Wheeler (1938) thought that Sinapis should 








Rhodora [Vol. 76 

properly be considered a part of Brassica. Sinapis 
arvensis of B&M cannot be transferred to Brassica 
because there is already a B. arvensis. Brassica kaber 
appears to be an older combination than B. will- 
denovii Boiss., as attributed to the Bahamas in 
Adams, 1972. 

Radicula brevipes —> Rorippa portoricensis var. pu- 
mila (O. E. Schulz) Stuckey. The name used in B&M 
and its synonyms are misapplied to the Bahamian 
taxon, according to Stuckey (1972). Stuckey inad- 
vertently maintained a neuter ending for the varietal 
epithet in his treatment; this is corrected above. 
Canara didyma — Coronopus didymus J. E. Smith. 
Coronopus (1757) is an older name than Canara 
(1792). Coronopus Zinn. (1757) is conserved over 
Coronopus Miller (1754), a genus of Plantaginaceae. 

Moringa moringa — Moringa oleifera Lam. In avoid- 
ing the tautonym, Moringa moringa, Ernst (1963) 
has picked up Moringa oleifera for the horseradish- 

Bryophyllum pinnatum — Kalanchoë pinnata (Lam.) 
Pers. Treatments of Kalanchoë and Bryophyllum 
(Baldwin, 1938; Hamet, 1907) have recognized 
Kalanchoë as the inclusive genus. 

Laurocerasus myrtifolia > Prunus myrtifolia  (L.) 
Urb. The cherry-laurel is best considered in the larger 
genus Prunus especially in view of other recent floras 
which have so placed it. 

Chrysobalanus pellocarpus ^ Chrysobalanus icaco L. 
Prance (1972) thought that there were insufficient 
means to separate C. pellocarpus from C. icaco and 
has treated them as taxonomic synonyms. 






Bahama Flora — Gillis 87 

Pithecellobium is conserved over Pithecolobium, and 
hence all species change to this orthography. The 
ram’s horn has been a source of confusion for a 
number of years. Pithecellobium keyense Britt. ex 
Coker in Shattuck was proposed to replace P. guada- 
lupense because the latter name was not applicable. 
Britton did not state why he considered the name 
inapplicable. In creating the new name, Britton 
(actually Coker) had no description and hence, 
created a nomen nudum. This situation was rectified 
in 1928 by Britton and Rose. Although the type of 
Mimosa guadalupensis Pers., the basionym, cannot 
be found at present in the Jussieu Herbarium at P, 
there appears to be no valid reason why this name 
cannot be taken up, pending proof that this plant 
does not represent the concept of the species known 
as “ram’s horn" in the West Indies and South Florida. 
Isely (1972) referred to specimens at the Delessert 
Herbarium and at Leiden which should be “critical,” 
but inasmuch as the basionym is linked by Persoon 
with “Hab ad Guadalupam (Herb. Juss.)," the lecto- 
type material should be sought in the Jussieu Her- 
barium. Why Britton and Rose indicated that this 
species was “not Inga guadalupensis Desv." (which 
is based on the same basionym) is unknown. The 
only Pithecellobium which occurs on Guadaloupe is 
this one. I am therefore retaining the name in B&M, 
but adding the parenthetical reference to the author 
of the basionym which was omitted in B&M: Pithe- 
cellobium guadalupense (Pers.) Chapm. 

The Lysiloma species remain as in B&M (Gillis and 
Stearn, in press), except for correcting gender to 

Anneslia haematostoma — Calliandra haematomma 
(Bert.) Benth. 

Anneslia formosa > Calliandra formosa (Kunth) 
Benth. Calliandra is conserved over Anneslia. The 









Rhodora [Vol. 76 

specific epithet haematostoma in B&M appears to be 
a misprint, judging from the basionym they cite. 
Despite efforts of Britton and Rose (1928) to frac- 
tionate Acacia into Bahamia and Lucaya, it seems 
best to maintain the three species of Acacia in B&M 
under the same names as in the flora: Acacia acuifera, 
A. choriophylla, and A. macracantha. 

Vachellia farnesiana — Acacia farnesiana (L.) Willd. 
Vachellia is best considered a generic synonym of 

Leucaena glauca — Leucaena latisiliqua (L.) Gillis et 
Stearn. Based on typification of an earlier name than 
either the epithet glauca or leucocephala (deWit, 
1961). Gillis and Stearn (in press) have shown that 
latisiliqua should be taken up. 

Acuan virgatum — Desmanthus virgatus (L.) Willd. 
var. virgatus. 

Acuan depressum —> Desmanthus virgatus var. de- 
pressus (Willd.) Turner. Desmanthus is conserved 
over Acuan. The two Bahama plants probably are 
best considered distinct only at the infraspecific 
level; therefore the treatment of Isely (1970) is fol- 

Neptunia — consult Windler (1966). 

Prosopis — consult Johnston (1962). 


Cassia. tora, —^ Cassia obtusifolia L. B&M suggest that 
Cassia tora and C. obtusifolia are synonyms and chose 
the former as the name to be used in the Bahama 
flora. Typification shows that such is not the case. 
DeWit (1955) selected a specimen in the Linnaean 
Herbarium as the type of C. tora L., but Brenan 
(1958) objected to this choice and selected a speci- 
men in the Hermann Herbarium (at BM) as the 
type. Based on the position of the petiolar gland, 
length of flowering and fruiting pedicels, anther 
shape, and presence of a 1.2-2 mm. areole on both 







Bahama Flora — Gillis 89 

sides of the seed, Brenan distinguished these two as 
distinct species. There is probably no Cassia tora 
in the New World according to Brenan. 
Chamaecrista chamaecrista — Cassia nictitans L. De- 
spite the efforts of Britton and Rose to fractionate 
Cassia into segregate genera Chamaecrista, Ditre- 
mexa, Peiranisia, ete., I maintain Cassia as an in- 
clusive genus. This necessitates different names for 
the six species placed in Chamaecrista in B&M. Cassia 
chamaecrista L. is based on a composite collection 
according to Pennell (1917). Therefore the correct 
name for the annual plant that is distributed through- 
out Eastern United States is generally accepted as 
C. fasciculata Michx. “Chamaecrista chamaecrista" 
of B&M should be rather the perennial species, Cassia 
nictitans L. I believe it is necessary to retypify all 
the names used in this complex in order to apply 
these names properly. 

Chamaecrista riparia > Cassia caymanensis C. D. 
Adams. Adams (1970) has shown that Cassia riparia 
is an illegitimate name, and therefore this plant must 
have a new epithet in Cassia. 

Chamaecrista lucayana — Cassia lucayana Britt. 
Chamaecrista caribaea — Cassia caribaea Northrop. 
Chamaecrista inaguensis > Cassia inaguensis Britt. 
Chamaecrista lineata > Cassia lineata Sw. These 
species revert to their original names in Cassia. 
Guilandina crista > Caesalpinia bonduc (L.) R. Br. 
Gray nickerbean has been shown by Dandy and Exell 
(1938) to be Caesalpinia (or Guilandina) bonduc. 
The epithet crista as used by Linnaeus refers to an 
Old World species. 

Guilandina bonduc — Caesalpinia divergens Urb. 
Guilandina ovalifolia — Caesalpinia ovalifolia Urb. 
Current thought among those working with this 
group of scrambling shrubs of the coastal regions of 
the West Indies is to treat them as a section of 
Caesalpinia, rather than a separate genus Guilandina. 






Rhodora [Vol. 76 

For some long period of time, G. bonduc was used to 
refer to the yellow-seeded nickerbean. As we have 
seen, proper typification by Dandy and Exell has 
shown that the name bonduc is misapplied when 
used thus. Among the synonyms given by B&M for 
this species, most apply to still other species. Caesal- 
pinia divergens seems to be the earliest name available 
for this species. The yellow-seeded nickerbeans in 
the Bahamas (with the exception of a new species 
to be described by Gillis and Proctor from Inagua) 
can be separated as follows: 

1. Seeds spherical, yellow; leaflets elliptic to sub- 
orbicular, obtuse or emarginate ............. 
Lee ee eee Caesalpinia ovalifolia (Urb.) Britt. 

1. Seeds oblong, bronze-brown; leaflets ovate, 
acute or acuminate ............. nee 

I accept B&M’s treatment of 1920 instead of the later 
treatment of Britton and Rose (1928) which recog- 
nized the segregate genus Vicarago. I see no reason 
to separate this taxon from Caesalpinia. 


Ateleia cubensis > Ateleia gummifera (Bert. ex DC.) 
Dietr. The epithet gummifera is an earlier epithet 
for this species (Rudd, 1968). 

Yakovlev (1967 a and b) has recognized a number of 
subspecific units in Sophora tomentosa L. For the 
moment, these seem difficult to accept as other than 
insular forms, and therefore I consider the taxon to 
be the single, variable S. tomentosa throughout the 
archipelago as in B&M. 

Cracca cinerea > Tephrosia cinerea (L.) Pers. 
Cracca cathartica — Tephrosia senna H.B.K. Although 
Cracca Benth. in Benth. et Oersted is conserved over 
Cracca L., Tephrosia Pers. is conserved over Cracca 
L., Needhamia Scopoli, and Reineria Moench. The 







Bahama Flora — Gillis 91 

correct generic name for the plants in the Bahamas 
is Tephrosia. Because the key to species in B&M is 
somewhat inadequate, a revised one follows: 

1. Leaflets 9-15; racemes opposite the leaves; pod 
2.5-4 mm. broad .. Tephrosia cinerea (L.) Pers. 
1. Leaflets 5-9; racemes terminal and opposite the 
leaves; pod 4-5 mm. broad ................. 
Tephrosia senna H.B.K. 

e «.4* 9» 9 6,8 he 6 6 8 à 5 € 8 9.0, 9 

Sesban sericea — Sesbania sericea (Willd.) DC. 
Sesban occidentalis —^ Sesbania emerus (Aubl.) Urb. 
Agati grandiflora — Sesbania grandiflora (L.) Pers. 
Sesbania is conserved over Sesban and Agati. The 
second of the species mentioned above was known 
only from fruit at the time of the writing of B&M. 
From collections made recently from the same popu- 
lation, I have determined it to be S. emerus. 
Stylosanthes — consult Mohlenbrock (1957). 
Meibomia supina —^ Desmodium canum (Gmel.) Schinz 
et Thell. 

Meibomia tortuosa —^ Desmodium tortuosum ( Mill.) 

Meibomia mollis ^À Desmodium glabrum (Mill.) DC. 
Desmodium is conserved over Meibomia. Supina 
cannot be used as an epithet for the first species above 
because it is based on a name which is a later 
homonym (Hedysarum supinum Sw. non H. supinum 
Chaix ex Villars). Incanum has often been taken 
up as a specific epithet for this plant also (based on 
Hedysarum incanum Sw.), but the basionym is a 
later homonym for H. incanum Thunb. Desmodium 
canum is based on the earliest available legitimate 
epithet. Desmodium glabrum is based on an earlier 
basionym than Meibomia mollis of B&M. 
Alysicarpus nummularifolius > Alysicarpus vaginalis 
(L.) DC. Alysicarpus vaginalis has been used for the 
false moneywort rather than a combination based on 
Hedysarum nummularifolium L. Schindler (1926) 








Rhodora [Vol. 76 

interprets the Linnaean material as being Indigofera 
echinata, in part, and Alysicarpus monilifer, in part, 
hence the choice of Hedysarum vaginale L. as the 
basionym rather than the earlier one, 
Ecastophyllum ecastophyllum > Dalbergia ecastophyl- 
lum (L.) Taub. Dalbergia’s having been conserved 
over Ecastophyllum eliminates the problem of the 
tautonym in B&M. 

Ichthyomethia | piscipula — Piscidia  piscipula  (L.) 
Sarg. Piscidia has been conserved over I[chthyome- 
thia. (See Rudd, 1969). 

Abrus abrus — Abrus precatorius L. The later epithet 
of Linnaeus is used to avoid the tautonym. 

Bradburya floridana — Centrosema floridanum (Britt.) 

Bradburya virginiana — Centrosema virginianum (L.) 
Benth. Bradburia has been conserved for a genus of 
Compositae over Bradburya of the Leguminosae. See 
also Lakela (1963). 

Canavali lineata — Canavalia rosea (Sw.) DC. 

Canavali bahamensis > Canavalia nitida (Cav.) Piper. 
Canavalia is conserved over Canavali. Sauer (1964) 
revised species of Canavalia, but apparently over- 
looked Johnston’s note (1949b) on publication of 
names referring to C. maritima. The basionym of 
this oft-used binomial has been treated as Dolichos 
maritima Aubl., but Thouars, in making the combina- 
tion Canavalia maritima, did not in fact cite the 
Aublet name; hence it is not a new combination, but 
a new name dating from 1813. Aublet’s name cannot 
then be transferred to Canavalia because it is pre- 
émpted by the Thouars combination, albeit for the 
same species. Alas, another name proposed in the in- 
terval is the oldest which must be used, Canavalia 
rosea (Sw.) DC. Species 2 of B&M (Canavali gladi- 
ata) is probably not in the Bahamas. All specimens 
bearing this determination by either Britton or Mills- 










Bahama Flora — Gillis 93 

paugh have been annotated by Sauer as C. maritima 
(=C. rosea). 

Cajan cajan —> Cajanus cajan (L.) Millsp. Cajanus 
is conserved over Cajan. The tautonym of B&M can 
thus be avoided. 

Dolicholus reticulatus ^ Rhynchosia reticulata (Sw.) 

Dolicholus swartzii > Rhynchosia swartzii (Vail) 

Dolicholus minima > Rhynchosia minima (L.) DC. 
Rhynchosia is conserved over Dolicholus. Dr. John 
Grear indicates (personal communication) that 
Rhynchosia caribaea based on Glycine caribaea Jacq. 
is not in the flora, thus agreeing with B&M. The 
latter is ironically not native to the New World, but 
is restricted to South Africa. 

Phaseolus lathyroides ^ Macroptilium lathyroides (L.) 
Urb. On the basis of having five calyx teeth rather 
than four, Macroptilium is segregated from the large 
genus Phaseolus. 

Dolichos lablab > Lablab purpureus L. Many workers 
(including Adams, 1972) use the generic segregate 
Lablab in preference to Dolichos. If one follows this 
trend, then the change above is indicated. Otherwise, 
Dolichos lablab may continue to be used. 

Dolichos insularis ^ Oxyrhynchus volubilis Brande- 
gee. This change follows studies by Rudd (1967). 
Vigna repens > Vigna luteola (Jacq.) Benth. in Mart. 
Although the epithet repens is the oldest name, it is 
invalidated in Vigna by V. repens Baker, published 
in 1876 (Merrill, 1910). 

Inoxalis intermedia > Oxalis intermedia A. Rich. 

Xanthoxalis corniculata > Oxalis corniculata L. Mod- 
ern treatment of Oxalidaceae would merge /noxalis 
and Xanthoxalis into Oxalis (Eiten, 1963). 









Rhodora [Vol. 76 

Cathartolinum  curtissii ^À Linum medium var. texa- 
num (Planch.) Fern. 
Cathartolinum corallicola —^ Linum bahamense var. 
corallicola (Small) Rogers. 
Cathartolinum bahamense > Linum bahamense North- 
rop var. bahamense. 
Cathartolinum bracei —^ Linum bahamense var. bracei 
(Small) Rogers. 
Cathartolinum lignosum — Linum bahamense North- 
rop. var. bahamense. Rogers (1963; 1968) has restored 
Linum as the correct name for the yellow-flowered 
species in the West Indies. I have followed his 1963 
treatment chiefly. 

Guaiacum — consult Porter (1972). 


Byrsonima cuneata — Byrsonima lucida (Mill.) DC. 
It is generally agreed that the plum-berry or locust- 
berry should be Byrsonima lucida, based on Malpighia 
lucida, an earlier epithet than cuneata. But upon 
whose M. lucida? Most authors attribute the name 
to Swartz, but his M. lucida is a later homonym for 
M. lucida Mill. Both names can be typified by material 
at BM. DeCandolle (1824) specifically indicated the 
Swartz material as basionym, but it is illegitimate as 
a later homonym. Kunth (1923) indicated that he 
thought that M. lucida should be a Byrsonima but did 
not make the transfer in accord with the Code. Rather 
than presume to make a new combination at this date 
myself, I interpret this situation in the light of his- 
tory. Swartz undoubtedly knew of Miller’s name and 
probably even examined herbarium material from the 
Chelsea Physic Garden upon which the lectotype of 
M. lucida Mill. (chosen by Gillis) has been selected. 
That he (Swartz) did not specifically indicate that 





Bahama Flora — Gillis 95 

he was taking up Miller's name is not important, it 
seems to me. There is ample precedent with Jacquin 
and Linnaeus, for instance, in assuming that the later 
author knew of the earlier work but did not, under 
existing convention, always cite the author of the 
earlier binomial, or place of publication. With this 
assumption, I shall follow Robertson (1971), in at- 
tributing the combination to DeCandolle with a lecto- 
type of Miller's name at BM. 

Cronquist's treatment of Alvaradoa (1944) would 
make the Bahamian populations Alvaradoa amorpho- 
ides subp. psilophylla (Urb.) Cronq. 

Elaphrium inaguense > Bursera inaguensis Britt. 
Elaphrium simaruba — Bursera simaruba (L.) Sarg. 
Bursera is conserved over Elaphrium. 

Badiera oblongata > Polygala oblongata — (Britt.) 
Blake. It seems desirable not to segregate the woody 
species of Polygala into the genus Badiera. 


Cicca disticha — Phyllanthus acidus (L.) Skeels. This 
name change follows Webster (1956-58). 
Margaritaria bahamensis > Margaritaria tetracocca 
(Baill) Webster. Webster (1956-58: J. Arnold 
Arbor. 38: 66) found that an earlier epithet applied 
to this species. 

Xylophylla epiphyllanthus —^ Phyllanthus epiphyllan- 
thus L. This treatment follows Webster (1967). 
Phyllanthus pruinosus > Phyllanthus  caroliniensis 
subsp. saxicola (Small) Webster. 

Phyllanthus pentaphyllus > Phyllanthus pentaphyllus 
C. Wright subsp. pentaphyllus. This treatment fol- 
lows Webster (1970). 













Rhodora [Vol. 76 

Croton flocculosus Geisl. > Croton flavens var. bal- 
samiferus (Jacq.) Muell.-Arg. It appears to me that 
Croton flocculosus is best treated as a variety of the 
widespread Croton flavens. It matches well the lecto- 
type of Croton balsamiferus Jacq. (lectotype from 
Martinique at BM), the basionym. 

Curcas curcas > Jatropha curcas L. Placement of the 
physie-nut in Jatropha seems to be the best place- 
ment taxonomically, and also avoids a tautonym. 
Adenoropiwm gossypifolium — Jatropha gossypifolia 
L. Adenoropium is best treated as a subdivision of 

Argythamnia — consult Ingram (1967). 

The place of publication of Acalypha alopecuroidea 
Jacq. should be Collect. 3: 196. 1790. 

Acalypha ostryaefolia > Acalypha ostryifolia Ridd. 
Pera bumeliaefolia > Pera bumeliifolia Griseb. These 
orthographic changes are required by the Seattle 

Manihot manihot > Manihot esculenta Crantz. This 
name is the earliest available to replace the tautonym 
used in B&M. 

Gymnanthes lucida — Ateramnus lucidus (Sw.) Rothm. 
Rothmaler (1944) and Dandy (1967) have resur- 
rected Ateramnus P. Browne to replace the later 
Gymnanthes Sw. Although I am of the opinion that 
Gymnanthes should be conserved over Ateramnus, 
conservation was already rejected once by the Special 
Committee for  Pteridophyta and Phanerogamae 
(Taxon 3: 241. 1954). 'There appears to be no 
choice but to adopt the new combination in 

Adenorima gymnonota > Euphorbia gymnonota Urb. 
Adenorima is treated as a section of Huphorbia by 
Webster (1967) and that treatment is followed here. 
Arthrothammus | cassythoides > Euphorbia cassytho- 
ides Boiss. Arthrothamnus differs too little from 
other forms of Euphorbia to be segregated from it. 















Bahama Flora — Gillis 97 

Tithymalus | trichotomus > Euphorbia trichotoma 
H.B.K. Tithymalus is treated by Webster (1967) as 
a section of Euphorbia. 

Aklema petiolaris > Euphorbia petiolaris Sims. 
Aklema differs too little from other forms of Eu- 
phorbia to be segregated from it. 

Chamaesyce wilsonit > Chamaesyce lecheoides var. 
wilsonii (Millsp.) Burch. 

Chamaesyce | lecheoides > Chamaesyce — lecheoides 
Millsp. var. lecheoides. 

Chamaesyce insulae-salis ^ Chamaesyce centunculo- 
ides (HBK.) Millsp. 

Chamaesyce exumensis —^ Chamaesyce lecheoides var. 
exumensis (Millsp.) Burch. These name changes fol- 
low the treatment by Burch (1966). 

Tricera bahamensis ^ Buxus bahamensis Baker in 
Hook. Tricera is best treated by the earlier name 
Buxus (Howard, 1962). 


Poison-ivy in the Bahamas is the typical subspecies: 
Toxicodendron radicans (L.) Kuntze subsp. radicans 
(see Gillis, 1971). 

Rhacoma coriacea, ^ Crossopetalum coriaceum North- 
Rhacoma — crossopetalum — Crossopetalum  rhacoma 
(Sw.) Hitchc. 
Rhacoma aquifolia + Crossopetalum aquifolium 
(Griseb.) Hitchc. 
Rhacoma ilicifolia > Crossopetalum ilicifolium (Poir.) 
Kuntze. Crossopetalum was revived by Brizicky 
(1964) as the earliest name for this genus. 
Although several recent rorkers have accepted Cas- 
sine as a widespread genus that would include 
Elaeodendron (Adams, 1972 and Ding Hou, 1963), 











Rhodora [Vol. 76 

I have chosen to follow Robson (1965) to recognize 
the segregate genus Elaeodendron for the Bahamas, 
thus leaving Cassine sensu stricto for a small African 


Thyana discolor > Thouinia discolor Griseb. Thou- 
inia Poit. is an earlier name than Thyana Hamilt. 
and is conserved over Thouinia Thunb. ex L.f. 
Allophylus cominia — Allophylus cobbe (L.) Raeusch. 
Leenhouts (1967) makes a strong case for recognizing 
only one worldwide species in this genus. He has 
found no morphological gaps which can be used to 
distinguish separate populations in any geographical 
sense. Having chosen to recognize only one species, 
Leenhouts chose A. cobbe as the all-encompassing 
name from two Linnaean names of the same date. 
Melicocca bijuga — Melicoccus bijugatus (L.) Jacq. 
The correct spelling of this name was worked out by 
Brizicky (1963). 

Sarcomphalus taylori — Ziziphus taylori (Britt.) 
M. C. Johnston. Johnston’s treatment (1964) merges 
Sarcomphalus with Ziziphus. 
Colubrina reclinata > Colubrina elliptica (Sw.) Briz. 
et Stern. 
Colubrina cubensis > Colubrina cubensis var. flori- 
dana M. C. Johnston. 
Colubrina colubrina > Colubrina arborescens (Mill.) 
Sarg. These changes follow the monographic treat- 
ment by Johnston (1971). 

Triumfetta — consult Ko Ko Lay, 1950. 

Phymosia — consult Fryxell (1971). 
Gayoides crispum — Herissantia crispa (L.) Brizicky. 








Bahama Flora — Gillis 99 

The earliest generic name available for this segre- 
gate of Abutilon is Herissantia (See Brizicky, 1968). 
Sida ciliaris — Sida ciliaris L. var. ciliaris. 

Sida carpinifolia —^ Sida acuta subsp. carpinifolia 
(L.f.) Borss. 

Sida, spinosa, — Sida spinosa var. angustifolia (Lam.) 

Sida hederaefolia > Sida javanensis Cav. emend. 
Borss. Recent treatments of Sida (Kearney, 1954b; 
Clement, 1957; Borssum Waalkes, 1966) do not all 
agree on treatment of the species in the Bahamas. I 
have followed Clement and Borssum Waalkes here. 
Malache scabra — Pavonia spicata Cav. var, spicata. 
Malache bahamensis > Pavonia bahamensis Hitchc. 
Pavonia is conserved over Malache. Kearney’s 
(1954a) treatment is followed here except for one 
point. He overlooked the fact that Pavonia scabra 
(B. Vogel) Juble et Quentin is a later homonym of 
P. scabra Presl. It must be replaced by P. spicata 
var. spicata. 

Malvaviscus sagraeanus > Malvaviscus arboreus var. 
mexicanus Schlecht. Schery (1942) has found an 
older name than the one given in B&M. 

Hibiscus bahamensis — Hibiscus brittonianus Kear- 
ney. Because H. bahamensis Britt. is a later homo- 
nym, Kearney (1954c) published a new name. 
Although a number of recent publications place okra 
in the genus Hibiscus as H. esculentus, Bates (1965) 
and Borssum Waalkes (1966) retained it in Abel- 
moschus, as in B&M, on the basis of the spathelike 
calyces which are basically adnate to the staminal 
tube and corolla, and which are circumscissilely de- 

Pariti tiliaceus — Hibiscus tiliaceus L. subsp. elatus 
(Hochr.) Borss. Borssum Waalkes (1966) is followed 
in his treatment of this taxon. 

Gossypium barbadense — Gossypium arborescens var. 
nadam (Watt.) Prockh. 


Rhodora [Vol. 76 

214 Gossypium punctatum > Gossypium hirsutum var. 


punctatum (Schum.) Hutch. Hutchinson (1943) is 
followed for the treatment of indigenous cotton 


The use of names in Melochia has been thoroughly 
considered by Goldberg (1967) who considered Mo- 
luchia and Melochia synonymous. The only changes 
from B&M therefore are orthographic, Because the 
keys in B&M are inadequate, a new key to the Bahama 
species is given here: 

1. Flowers in sessile glomerules, many per cluster; 
sinus between calyx teeth acute; fruit a 5-parted 
coccus less than 3.5 mm. in diameter ........ 
SNMP Melochia nodiflora Sw. 

. Flowers pedicelled, 3-10 per cluster; sinus be- 
tween calyx teeth rounded to truncate; fruit 
a 5-angled capsule, greater than 5 mm. in 
diameter .......... ee RII 2. 
2. Capsule with rounded angles and unbranched 

hairs; inflorescences mostly in axillary cymes 
or appearing terminal; foliage leaves 
glabrous or with scattered, simple hairs; 
leaves only slightly lighter in color on lower 
surface than on upper ........... enne 
IM Melochia pyramidata L. 
Capsule sharply angled, with dense stellate 
hairs; inflorescences mostly opposite the 
leaves, never terminal; foliage leaves densely 
stellate; leaves tending toward being dis- 
colorous, much lighter on the lower surface 
than upper due to dense pubescence ...... 
2.3232 *2»*«***3** Melochia tomentosa L. 

278 Waltheria americana —^ Waltheria indica L. An ex- 

amination of Linnaean material and resultant lecto- 
typification has resolved the question of a name for 





Bahama Flora — Gillis 101 

the common Waltheria in accord with Brizicky (1966) 
and the example used in the International Code (Art. 
57). The lectotype (chosen by Gillis) of Waltheria 
indica L. is in the Hermann Herbarium at BM. The 
lectotype of Waltheria americana L. is at LINN 
(Savage Catalog No. 852: 1). They represent the 
same taxon; both names originate from the same 
date (1753). Robert Brown (in Tuckey, Narr. Exp. 
River Zaire 484. 1818) appears to be the first to 
adopt the name W. indica L. for the combined species, 
and therefore should be followed. 

The descriptions and separation of the two species 
of Waltheria in B&M are ambiguous. The foliage of 
W. bahamensis is decidedly bronze in the field; her- 
barium specimens do not show this character well, 
but it is useful as a field character. The following 
key should assist in making determinations: 

Foliage green; branches of stellate hairs 2-4 mm. 
Ing 550500 Lot Waltheria indica L. 
Foliage bronze; pubescence in small tufts, branches 
of stellate hairs less than 1 MIN ...... ers 
Waltheria bahamensis Britt. 

Ayenia pusilla > Ayenia insulicola Cristobal. Except 
for an orthographic change necessitated by the Seattle 
Code, the Ayenia species should be named according 
to Cristobal (1960). 

Ascyrum linifolium > Hypericum hypericoides (L.) 
Crantz. The nomenclature of Ascyrum has under- 
gone much discussion (Adams, 1957; Adams and 
Robson, 1961). The treatment of Adams and Robson 
(ibid.) has been followed here, 


Myroxylon ilicifolium > Xylosma ilicifolia Northrop. 
Myroxylon bahamense —^ Xylosma bahamensis (Britt.) 
Standl. Xylosma G. Forst. has been conserved over 






Rhodora [Vol. 76 

Myroxylon J. R. et G. Forst. as a genus of Flacourtia- 
ceae (Myroxylon L. has been conserved in the Legu- 
minosae). The question of whether these two species 
are, in fact, distinct still needs to be explored further. 

Passiflora pallida > Passiflora suberosa L. The two 
names above are synonyms of the same date; they 
were united by Killip (1938). See also Brizicky, 1961. 
Passiflora species in the Bahamas will be evaluated 
in a separate paper. 

Mentzelia — consult Darlington (1934). 

Parsonsia parsonsia — Cuphea parsonsia (L.) R. Br. 
Parsonsia is conserved for a genus of Apocynaceae; 
it therefore cannot be used for a genus of Lythraceae. 

Conocarpus erecta > Conocarpus erectus L. The 
generic name Conocarpus should be treated as mascu- 
line in accordance with the International Code 
(Recommendation 75A). In Stearn (1966) the sub- 
stantive termination -carpus is consistently treated 
as masculine. Stearn (personal communication) sug- 
gests that Linnaeus may have treated Conocarpus as 
feminine by analogy with many classical second- 
declension names of trees. 


Eugenia buxifolia > Eugenia foetida Pers. The 
Myrtaceae need to be reworked in the light of Mc- 
Vaugh’s generic guidelines and definitions (1956; 
1968). The change indicated above is made in accord 
with his analysis of types (McVaugh, 1973). 
Pimenta pimenta — Pimenta dioica Merr. This change 
is made to avoid a tautonym. 










Bahama Flora — Gillis 103 

Anamomis longipes > Psidium longipes (Berg) Mc- 
Vaugh var. longipes. 

Anamomis bahamensis — Psidium longipes var. orbi- 
culare (Berg) McVaugh. 

Anamomis lucayana — Myrcianthes fragrans (Sw.) 
McVaugh. The first two of these changes is in accord 
with the interpretation of these species by MeVaugh 
(1973). Psidium longipes var. orbiculare is further 
elaborated in the same paper by McVaugh by de- 
scribing unnamed races which can be distinguished. 
Myrcianthes fragrans is defined in an earlier paper 
by MeVaugh (1963). 


Jussiaea suffruticosa > Ludwigia octovalvis subsp. 
sessiliflora (Micheli) Raven. This name change fol- 
lows the treatments of Ludwigia by Raven (1962; 


Foeniculum foeniculum — Foeniculum vulgare Gaertn. 
This change is made to avoid a tautonym. 


Icacorea paniculata — Ardisia escallonioides Cham. 
et Schl. 

Icacorea guadalupensis > Ardisia obovata Desv. 
Ardisia is conserved over Icacorea. The Bahama 
plants need further study and interpretation, but they 
appear to be named best as stated above. 

Rapanea guianensis > Myrsine floridana A. DC. 
Rapanea guianensis Aubl. may be typified by a speci- 
men of Aublet's in P (lectotype of Gillis). It repre- 
sents a plant with lustrous, large leaves from northern 
South America. The northern West Indian and Flo- 
ridian species has dull, smaller leaves. Futhermore, 
it represents the group of species with a globose 
stigma, a character which has been used to separate 
Myrsine from Rapanea. Stearn (1969) made the 








Rhodora [Vol. 76 

combination Myrsine punctata (Lam.) Stearn for this 
species, but the name is a later homonym for a species 
of the Pacifie region which had been published several 
years earlier. The earliest available name for this 
plant is Myrsine floridana A. DC. If one prefers 
this species in Rapanea, then its name should be 
R. punctata (Lam.) Lundell. 


Samolus floribundus — Samolus parviflorus Raf. This 
change is made in aecordance with Channell and Wood 
(1959) ; it is based on an earlier epithet. 


Chrysophyllum — consult Cronquist (1945). 
Sideroxylon foetidissimum — Mastichodendron foeti- 
dissimum (Jacq.) Cronq. This change is made in ac- 
cord with the treatment by Cronquist (1946a). 
Lucuma serpentaria > Pouteria domingensis (Gaertn.) 
Cronq. var. domingensis. This change follows the 
treatment by Cronquist (19462). 

Dipholis salicifolia > Bumelia salicifolia (L.) Sw. 
Although Cronquist (1945) retained Dipholis as a 
segregate genus, Stearn (1968) merged it with 
Bumelia. It is the latter treatment which is followed 

Bumelia angustifolia > Bumelia celastrina H.B.K. 
Bumelia loranthifolia > Bumelia americana (Mill.) 
Stearn subsp. americana. 

Bumelia bahamensis > Bumelia americana (Mill.) 
Stearn subsp. americana. The Bumelia complex in 
the West Indies has been treated by Stearn (1968), 
a treatment followed here. The following key should 
help clarify the species in the Bahamas: 

1. Ovary glabrous; endosperm copious; leaves 
longer than 7 cm.; terminal shoots never be- 
coming thorns .. Bumelia salicifolia (L.) Sw. 

1. Ovary pubescent; endosperm little or none; 

1974] Bahama Flora — Gillis 105 

leaves shorter than 6 em. ; terminal shoots often 
thorny a C ok oie CE ELE 2. 
2. Leaves narrow, linear to spathulate, 2-10 
mm. broad (juveniles may be larger) ; fruit 
oblong; plant inhabiting shores, mangal, 
and SUNG areas. va ck ee ts re 

2. Leaves obovate to oblanceolate, 1-4 mm. 
broad; fruits globose; inhabiting uplands .. 
A cR Bumelia americana, (Mill.) Stearn 
subsp. americana. 

324 Mimusops emarginata — Manilkara bahamensis 
(Baker) Lam. et Meeuse. Probably no species in the 
Bahamas is easier to identify and yet harder to name 
than the wild dilly! In addition to the long list of 
synonyms given in B&M, it has been called Achras 
emarginata (L.) Little, Mimusops jamaiqui (Wright) 
Dubard, M. jamaiqui subsp. emarginata, (L.) Cronq., 
and Manilkara parvifolia (Nutt.) Dubard, to name 
a few. For the moment, I have selected Manilkara 
bahamensis (Baker) Lam. et Meeuse. Thus, it re- 
sides in the same genus preferred by Moore and 
Stearn (1967) for its close relative, the sapodilla, 
and also preferred by Cronquist (1945). 

324 Sapota achras — Manilkara zapota (L.) P. van Royen. 
By the same token as with the wild dilly (above), the 
cultivated sapodilla has had a variety of names, 
many of which are permutations of each other. Moore 
and Stearn (ibid.) analyzed the problem thoroughly 
and leave a choice of names depending upon the 
breadth of one's generic concepts. Along with them, 
I choose the one indicated above. 

325 Maba crassinervis > Diospyros crassinervis (Krug 
et Urb.) Standl. Maba is treated as being a synonym 
of Diospyros. The name given above in Diospyros is 
temporary. A new combination will soon be pub- 
lished by Mr. Frank White (Oxford). 









Rhodora [Vol. 76 


The Mayepea bumelioides of B&M will be placed in 
Chionanthus by Stearn at a later date, at which time 
a new combination will be published. 

Based on Shinners’s typification (1957), the name of 
the marsh gentian remains the same, but the author 
of the binomial and place and date of publication 
change from what is given in B&M: Eustoma exalta- 
tum (L.) Salisb., Parad. Lond. t. 34. 1806. 
Sabbatia campanulata — Sabatia stellaris Pursh. 
Sabbatia simulata — Sabatia stellaris Pursh, Wilbur 
(1955) and Perry (1971) have studied the taxonomy 
of Sabatia in detail. As a consequence, one species 
only is considered to be in the Bahamas. The correct 
spelling of the generic name should also be noted. 

Nymphoides aureum — Nymphoides grayana (Griseb.) 
Kuntze. This name change is made in accord with 
the treatment by Ornduff (1969). It should be noted 
that this so-called endemic Nymphoides (according 
to B&M) is also found in Cuba. 

Plumiera obtusa — Plumeria obtusa L. var. obtusa. 
Plumiera inaguensis > Plumeria obtusa L. var. ob- 
Plumiera bahamensis —^ Plumeria obtusa L. var. ob- 
Plumiera rubra — Plumeria rubra L. 
Plumiera sericifolia, + Plumeria obtusa var. sericifolia 
(C. Wright) Woodson, Woodson (1938) reworked 
the species of Plumeria, taking up the correct spelling 
of the generic name. He considered several of the 
"species" of B&M to be insular races of P. obtusa 
var. obtusa. 








Bahama Flora — Gillis 107 

Woodson (1936) recognized Neobracea bahamensis 
as endemic, but also noted that there are three other 
species in Cuba, i.e, it is not a monotypic genus as 
stated in B&M. 

Stearn (1964) confirmed the name of the periwinkle 
as Catharanthus roseus (L.) G. Don, 

Echites echites > Echites umbellata Jacq. var. um- 

Rhabdadenia paludosa > Rhabdadenia biflora (Jacq.) 

Rhabdadenia sagraei —^ Angadenia berteri (A. DC.) 

Urechites lutea — Urechites lutea var. serica Long. 
Woodson (1936) redefined the generic limits of a 
number of American Apocynaceous plants. His treat- 
ment is followed here. Long (1970a), in studying 
populations in South Florida, treated Urechites such 
that the Bahamian populations are recognized as a 
distinct pubescent population, U. lutea var. sericea. 
Because the Apocynaceous vines are not adequately 
keyed in B&M, and because the names of all of 
them have been changed, the following key has been 
prepared for use in separating them: 


1. Corolla white or ivory, usually more than 5 cm. 
long; leaves remaining flat or folding along the 
midrib when pressed ; follicles of pair divergent, 
not tending to curve back like pincers, 4-10 mm. 
ihiek sou ily doe rer EDITI 2. 
2. Follicles of a pair widely divergent (greater 

than 180?) ; tube of corolla salverform with 
rotate lobes; cylindric part of corolla 10-15 
times the length of the sepals; calyx lobes 
free nearly to base, linear to linear-lanceolate 

. Echites umbellata Jacq. var. umbellata. 

2. Follicles of a pair divergent less than 90°; 
tube of corolla funnelform; cylindric part of 




Rhodora [Vol. 76 

corola 2-3 times the length of the sepals; 
calyx lobes free scarcely more than 2/3 the 
length, ovate, to 2.5 mm. broad .......... 

.. Rhabdadenia biflora (Jacq.) Muell.-Arg. 

1. Corolla yellow, usually less than 5 cm. long; 
leaves tending to curl under at the edges when 
pressed ; follicles of a pair diverging at an angle 
of less than 60°, curving back to touch near their 
tips, resembling a pair of pincers, 1.5-3 mm. 
thick .............eleele eh 3. 
3. Vine scarcely 1 m. long; corolla 2-4 cm. 
long; leaves oblong, usually less than 1 cm. 
broad; pedicels glabrous; calyx lobes free 
less than half their length, 1-1.5 mm., acumi- 
nate; cylindric part of corolla visible most 

of its length (i.e, not hidden by sepals); 
petiole 2 mm. long ..................... 
TEE Angadenia berteri (A.DC.) Miers 

3. Vine up to 3.5 mm. long; corolla 4-5 cm. 
long; leaves obovate or elliptic to 3 cm. 
broad; pedicels densely pubescent; calyx 
lobes free nearly to base, 8-12 mm. long, 
linear; cylindric portion of corolla hidden 
by sepals; petiole 1 em. long ............. 
ZEE Urechites lutea var. sericea Long 

Rauwolfia ` tetraphylla > Rauvolfia nitida Jacq. Rao 
(1956) thought that Rauvolfia tetraphylla is indige- 
nous to the Greater Antilles, Central America, and 
South America as far as Peru and Venezuela, The 
Bahamian species is R. nitida, which is not a synonym 
of R. tetraphylla as suggested in B&M. 

Vallesia glabra — Vallesia antillana Woodson. Wood- 
son (1937, 1938b) found that the true Vallesia glabra 
is a plant of Mexico and the Pacific Coast of South 
America. The species of southern peninsular Florida 
and the West Indies that has corollas twice the length 
of those of V. glabra needed a name. The name V. 
glabra of B&M is thus misapplied. 











Bahama Flora — Gillis 109 


Metastelma | northropiae > Cynanchum northropiae 
(Schltr.) Alain, 

Metastelma hamatum > Cynanchum caribaeum Alain. 
Metastelma inaguense > Cynanchum inaguense (Vail) 
Howard et Dunbar. 

Metastelma linearifolium > Cynanchum savannarum 

Metastelma eggersii > Cynanchum eggersii (Schltr.) 

Metastelma palustre > Cynanchum angustifolium 
Pers. Metastelma species are considered synonymous 
with Cynanchum. (See Woodson, 1941; Alain, 1955). 
Much work needs to be done on the Bahamian species, 
but almost certainly several will be united. I shall 
refrain from making transfers to Cynanchum of 
several names which have not already been trans- 
ferred because I feel certain at this point that these 
names are synonyms of other names in the flora. 
(See also Howard and Dunbar, 1964). Merrill and 
Hu (1949) discovered that Muhlenberg had found 
Cynanchum angustifolium to be the eariest legitimate 
name for the last species listed above. They indicated 
the extensive synonyms for this species, a few of 
which had never ben included in the standard indices. 
Philibertella clausa > Sarcostemma clausum (Jacq.) 
R. & S. Sarcostemma is an earlier name than Phili- 
bertella (Woodson, 1941). 


Evolvulus glaber —^ Evolvulus convolvuloides ( Willd.) 

Evolvulus alsinoides > Evolvulus alsinoides var. gri- 
sebachianus Meissn. in Mart. 

Evolvulus linifolius > Evolvulus alsinoides var. lini- 
folius (L.) Baker. Stearn (1972) has modified the 
nomenclature of several species of Evolvulus in the 
flora in the course of typification of their names. The 





Rhodora [Vol. 76 

last taxon listed above is now believed not to occur 
in the flora. 

Jacquemontia jamaicensis ^ Jacquemontia havanensis 
(Jacq.) Urb. Dr. Kenneth Robertson (personal com- 
munication) has pointed out that this name change 
is required because of an earlier basionym. 
Calonyction aculeatum — Ipomoea alba L. 
Calonyction tuba — Ipomoea macrantha R. et S. Both 
Ooststroom (1940) and Gunn (1972) have worked 
with Calonyction. Gunn’s nomenclature is followed 

Quamoclit quamoclit ^ Ipomoea quamoclit L. 
Quamoclit coccinea > Ipomoea hederifolia L. Quamo- 
clit, like Calonyction, is treated as a section of Ipo- 
moea. (see Ooststroom, 1953). 

Ipomoea cathartica — Ipomoea acuminata (Vahl) R. 
et S. 

Ipomoea pes-caprae > Ipomoea pes-caprae subsp. bra- 
Siliensis (L.) Ooststr. This treatment of Ipomoea 
follows that of Ooststroom (1940) and St. John 
(1970). The citation for Ipomoea stolonifera in B&M 
is incorrect; there is an earlier publication date of the 
combination: Ipomoea stolonifera (Cyrill.) Gmel., 
Syst. Veg. I: 345. 1796. 

Ipomoea dissecta + Merremia dissecta (Jacq.) Hall. f. 
Merremia is segregated from Ipomoea on the basis of 
its smooth pollen and its corolla without a red or 
purple eye (O'Donell, 1941; Ooststroom with Hoog- 
land, 1953; Verdcourt, 1963). 

Dichondra. Consult Tharp and Johnston (1961). 

Cuscuta pentagona — Cuscuta campestris Yuncker. 
This species needed a new name because of confusion 
in the old one (Yuncker, 1932). 
Marilaunidium jamaicense > Nama jamaicensis L. 











Bahama Flora — Gillis 111 

Nama is an earlier name than Marilaunidium. Fur- 
thermore, Nama jamaicensis L. is the conserved type 


Sebesten sebestena — Cordia sebestena L. 

Varronia globosa > Cordia globosa var. humilis 
(Jaeq.) Johnston. 

Varronia bahamensis > Cordia bahamensis Urb. 
Varronia brittonii > Cordia brittonii (Millsp.) Mac- 

Varronia lucayana — Cordia lucayana (Millsp.) Mac- 
bride. Sebesten and Varronia should be treated as 
synonyms of Cordia (Macbride, 1916; Johnston, 

Rochefortia bahamensis — Rochefortia spinosa (Jacq.) 
Urb. According to Lefor (1968), the Rochefortia in 
the Bahamas is not an endemic species, but the same 
as that found in Cuba. 

There is debate over the generic selection for bay 
lavender. For the moment, I maintain Mallotonia, 
as in several other recent floras (Gooding, Loveless, 
and Proctor, 1965; Adams, 1972). 

Heliotropium parviflorum — Heliotropium angiosper- 
mum Murray. The change of name is required because 
H. angiospermum is an earlier name. 

Heliotropium inundatum — Heliotropium procumbens 
Mill. Heliotropium procumbens is an earier name for 
the same species. Both names can be typified by 
specimens at BM; the lectotypes have been selected 
by me. 


Valerianoides fruticosa > Stachytarpheta fruticosa 
(Millsp.) B. L. Robinson. 

Valerianoides jamaicensis —> Stachytarpheta jamai- 
censis (L.) Vahl. Stachytarpheta is conserved over 








Rhodora [Vol. 76 

Bouchea prismatica > Bouchea prismatica var. longi- 
rostra Grenzen. This treatment follows the revision 
of Bouchea by Grenzenbach (1926). Moldenke (1971) 
believes that the typical variety is also present in 
the Bahamas. 

Priva — consult Kobuski (1926). 

Lippia stoechadifolia > Phyla stoechadifolia — (L.) 

Lippia nodiflora — Phyla nodiflora (L.) Greene var. 

Lippia reptans — Phyla nodiflora var. reptans (H.B.K.) 
Moldenke. These changes are made in accord with 
the treatment of Phyla by Moldenke (1965). Lippia 
geminata of B&M is probably not found within the 
archipelago. The appropriate treatment of this bi- 
nomial would be Lantana microcephala A. Rich. in 
any event. 

According to treatment by Long (1970a) our variety 
of Lantana ovatifolia would be the typical variety 
(var. ovatifolia). 

Using Moldenke's treatments (1958a, b, and c) of 
Citharexylum, I have prepared the following key to 
distinguish the Bahamian species: 

Pedicels less than 1.2 mm. long during anthesis; 
pyremes 2-loculate, fruit not shiny; leaves + pilose, 
closely reticulate-veined, the reticulation prominent 
above and below ... Citharexylum fruticosum L. 

Pedicels 2 mm. long or more during anthesis; 
pyremes 1-loculate, fruit shiny; leaves glabrous, 
widely reticulate-veined, reticulation obscure above 
Citharexylum caudatum L. 

Moldenke (1958a; 1971) has considered 5 infra- 
specific taxa of C. fruticosum to exist in the Bahamas: 
var. fruticosum, var. smallii Moldenke, var. subvil- 
losum Moldenke, var. villosum (Jaeq.) O. E. Schultz, 
and forma bahamense Moldenke, Having made field 







Bahama Flora — Gillis 113 

studies on the characters upon which these taxa are 
based, I consider these to be insular forms or popula- 
tions without names. For those who wish to name 
these variations, an appropriate key is found in 
Moldenke (1958a). 

Callicarpa — consult Moldenke (1936). 

Petitia — consult Moldenke (1937). 

Volkameria aculeata —^ Clerodendrum aculeatum (L.) 
Schlecht. var. aculeatum. Volkameria is treated by 
Moldenke (1971) as a section of Clerodendrum. The 
place of citation for C. aculeatum is incorrect in 
B&M. The author of the combination is Schlechten- 
dahl, and the citation is Linnaea 6: 750. 1831. 
Clerodendrum fragrans > Clerodendrum philippinum 
Schauer. Howard and Powell (1968) found that C. 
fragrans was an illegitimate name. Moldenke (1971) 
considers the Bahamian population to be var. pleni- 
florum Schauer. 

Avicennia nitida — Avicennia germinans (L.) L. 
Stearn (1958) found this earlier epithet which ap- 
plies to the species of black mangrove in the West 
Indies, but failed to note that its earliest combination 
was by Linnaeus himself. 


Melosmon cubense — Teucrium cubense Jacq. Epling 
(1925) united Melosmon and Teucrium. 

Leonurus — consult Epling (1925). 

The place of publication for Leonotis nepetifolia (L.) 
R.Br. is the Prodromus florae Novae Hollandiae, 
p. 504. 1810, and not as given in B&M (Epling, 

In Epling’s treatment (1938-39), Salvia serotina L. 
and S. micrantha Vahl (indicated as synonyms in 
B&M) were separated on rather minor characters. 
Through the courtesy of Dr. Mildred Mathias and 
Dr. Raymond Harley, I have seen an unpublished 
manuscript of Epling and Carlos Jativa in which 




Rhodora [Vol. 76 

these species were reunited. It seems best to continue 
to treat them as synonymous. (Permission of Carlos 
Jativa to refer to this unpublished material is ac- 
knowledged with appreciation.) 

On the basis of rather minor characters of the calyx, 
Shinners (1962) reworked the Micromeria brownei 
complex, recognizing the Bahamian populations as 
M. bahamensis Shinners. Because my own collections 
from Inagua more closely resemble the taxon which 
Shinners cites from Hispaniola, I believe more work 
needs to be done before the matter is resolved. Chiefly 
because of the minor nature of the characters which 
are used in defining Shinners's species, I continue to 
recognize a broadly treated M. brownei (Sw.) Benth. 


Physalis angulata > Physalis angulata L. var. angu- 

Physalis turbinata — Physalis cordata Mill. 
Physalis pubescens > Physalis barbadensis Jacq. 
These changes are in accord with the treatment by 
Waterfall (1967). 

Solanum verbascifoltum — Solanum erianthum D. 
Don. Roe (1968) found that the old name for this 
plant did not apply to the species in our flora. 
Lycium spathulifolium > Lycium tweedianum var. 
chrysocarpum (Urb. et Ekm.) Hitche. In the mono- 
graph of Lycium (Hitchcock, 1932), the Bahamian 
plant required a name change. Although it is possible 
that the species of the Florida Keys (L. carolinianum 
Walt.) is in the Bahamas, it has yet to be found. 

Maurandya antirrhinaeflora > Maurandya antirrhini- 
flora H. et B. ex Willd. This orthographic change is 
required by the Seattle Code. I can now report this 
species to be on the far side of the archipelago from 
New Providence as recorded in B&M: Gillis 11780 
from Grand Turk. 









Bahama Flora — Gillis 115 

Bramia monnieri > Bacopa monnieri (L.) Pennell. 
Bacopa has been conserved over Brami and Moniera. 
It is an earlier name than Bramia. 

Although Pennell (1935) segregated Erinus from 
Mecardonia, I believe that Mecardonia is best treated 
as an inclusive genus, thus maintaining Mecardonia 
procumbens (Mill.) Small as in B&M. 

Afzelia cassioides > Seymeria  cassioides (Walt.) 
Blake. Seymeria is conserved over Afzelia. 
Buchnera elongata —> Buchnera floridana Gandoger. 
Philcox (1965) thinks that B. elongata is a mis- 
applied name for the Florida and Bahama popula- 

Setiscapella subulata — Utricularia subulata L. 
Stomoisia cornuta — Utricularia cornuta Michx. Tay- 

lor (1967) includes Setiscapella and Stomoisia within 


Macrocatalpa punctata — Catalpa punctata Griseb. 
var. punctata. This change is made in accord with 
the treatment of Catalpa by Paclt (1952). 


Sesamum orientale —^ Sesamum indicum L. These 
names are synonymous binomials of the same publica- 
tion date. Sesamum indicum L. has been used more 
commonly than the other name. Hill (1939) pointed 
out that DeCandolle (Pl. Rar. Jard. Genève 18, t.5 — 
1889) was the first to unite the two names under 
Sesamum indicum. 


Blechum brownei should be retained despite the at- 
tempt of some to change the name of this species to 
Blechum pyramidatum. The Linnaean binomial is 
Ruellia bltechum, Lamarck renamed the species Bar- 










Rhodora [Vol. 76 

laria pyramidata, but this name is illegitimate, being 
superfluous when published. Jussieu later published a 
new name in Blechum; this is accepted because the 
tautonym which would be created by transferring the 
Linnaean epithet is illegitimate (Bremekamp, 1938; 
Long, 1970b). 

Gerardia droseroides > Stenandrium droseroides Nees 
in DC. 

Gerardia bracteosa — Stenandrium bracteosum (Britt. 
et Millsp. Britt. ex Leonard). Stenandrium is con- 
served over Gerardia. 

Anthacanthus spinosus — Oplonia spinosa (Jacq.) Raf. 
Stearn (1971) has studied the generic complex of 
which this species is a part and has recognized our 
plant to be an Oplonia. 

Diapedium assurgens  Dicliptera assurgens  (L.) 
Juss. Dicliptera is conserved over Diapedium (see 
also Long, 1970b). I have chosen the lectotype to be 
at LINN (No. 28:23 according to the Savage Catalog, 


The Rubiaceae are in need of in-depth treatment as 
a whole. Names accepted here may be subject to re- 
vision as more work is done. 

Oldenlandia callitrichoides > Hedyotis callitrichoides 
(Griseb.) Lewis. Merrill and Metcalf (1942) and 
Lewis (1961) have united Oldenlandia and Houstonia 
with Hedyotis. Lewis’s treatment is followed here. 
Rachicallis americana — Rhachicallis americana (Jacq.) 
Hitche. The original spelling of the generic name has 
been restored. Despite the fact that it has been 
written without the first “h” ever since the original 
publieation, there seems no reason not to accept the 
original orthography considering the Greek origin. 
Nickerson and Tripp (1973) have reported this species 
to have either male or perfect flowers, never female. 
Randia mitis — Randia aculeata L. These two names, 







Bahama Flora — Gillis 117 

synonyms of the same date of publication, have been 
united under R. aculeata. 

Catesbaea, parviflora — Catesbaea parviflora var. sep- 
tentrionalis Krug et Urb. ex Urb. The Bahamian 
populations fit this variety better than do the typical 
species form. 

Hamelia erecta > Hamelia patens Jacq. These names 
are synonyms of the same date of publication. Thomas 
Elias (personal communication) has informed me 
that L’Héritier was the first to unite them, and should 
be followed. 

Guettarda taylori —^ Guettarda nashii Britt. et Millsp. 
Guettarda inaguensis > Guettarda nashii Britt. et 
Millsp. There does not seem to be sufficient difference 
to maintain three distinct species of Guettarda sup- 
posedly endemic to the vicinity of Matthew Town, 
Inagua. Having studied both the type specimens (at 
NY) and material in the field, I have concluded that 
Britton and Millspaugh named specimens, not species. 
Guettarda inaguensis is represented by a type that 
appears to have come from an aberrant form with 
long lengths of bare branches and leaves only near 
the tips. Guettarda taylori is represented by a type 
that has no flowers left and for which no fruits were 
ever known. Because of the inadequancy of these two 
specimens, it seems best to select the name for the 
united species by the type possessing the most ade- 
quate material. Inasmuch as these three species 
names originate from the same data, I am choosing 
G. nashii as the name to be used when all three are 
united. The populations of Guettarda in the vicinity 
of Matthew Town, although variable, appear to me 
to be best treated as a single species. 

Stenostomum lucidum — Antirhea lucida (Sw.) Hook. 
f. in Benth. et Hook. f. 

Stenostomum  myrtifolium — Antirhea myrtifolia 
(Griseb.) Urb. 

118 Rhodora [Vol. 76 

415 Stenostomum densiflorum — Terebraria resinosa 
(Vahl) Sprague. Stenostomum as used by B&M is 
better divided into Antirhea and Terebraria. Antirhea 
is an older name than Stenostomum and hence must 
be the name used for those species traditionally placed 
in Stenostomum. The tortuous history of the name 
of Terebraria resinosa is recounted thoroughly by 
Sprague (1932). Sprague recognized two West In- 
dian species of Terebraria, differing only in shape 
of the areoles of the leaf venation. In his treatment, 
the Bahamian population would be T. densiflora. I 
consider these differences to represent only insular 
forms, not worthy of nomenclatural separation. A 
key to the species in the Bahamas of the old Stenos- 
tomum follows: 

1. Ovary 2 (-4) loculate; flowers without bracts; 
anthers half-exserted; stipules deciduous; foli- 
age without resinous secretions .... Antirhea 
lucida (Sw.) Hook. f. in Benth. et Hook. f. 

1. Ovary 4-6 loculate; flowers subtended by minute 
bracts; anthers included; stipules persistent; 
foliage resinous-viscid ................0-. 2. 

2. Leaves 3 cm. long or less, elliptic to obovate- 
oblong; diffusely-branched shrub with leaves 
crowded near ends of branches; midvein very 
pronounced, often white; 2-4 flowers per 
branch of inflorescence; stipules ovate, not 
forming a collar around stem; inflorescences 
paired on either side of stem, but not 
branched ................ eee 

2. Leaves 4-9 cm. long, oblong-lanceolate ; 
coarsely-branched shrub or small tree with 
leaves widely scattered; midvein only slightly 
more pronounced than lateral veins; more 
than 6 flowers per branch of inflorescence; 
stipules forming a collar 2-4 mm. broad 




Bahama Flora — Gillis 119 

around stem; inflorescence bifid with flowers 
only on inside (adaxial surface) of branches 
Terebraria resinosa (Vahl) Sprague. 

Psychotria undata > Psychotria nervosa Sw. Jacquin 
described P. undata in the Plantarum rariorum horti 
caesarei schoenbrunnensis. According to Stafleu 
(1967), the title page date of 1798 is probably in 
error, and actual date of publication may have been 
as late as 1803. In any event, Swartz in his Prodro- 
mus (1788) published the name Psychotria nervosa 
for the same species, clearly the earliest date of pub- 
lication. Psychotria nervosa Benth, (1841) and P. 
nervosa D. Don (1825) are later homonyms. 

Ernodea cokeri > Ernodea taylori Britton. 

Ernodea nashii o Ernodea millspaughii Britton. It 
is difficult to make hard-and-fast decisions on the 
Ernodea problems in the Bahamas at this juncture. 
Whether E. littoralis and E. angusta are truly distinct 
is questionable. Long (1970a) has treated E. angusta 
as a variety of E. littoralis which may be the best 
way to handle these perplexing taxa. Until intensive 
study is made of these two species with comparisons 
to the other Ernodea taxa in the Bahamas, I shall 
continue to recognize these two as species. Distinc- 
tions among the other four species do not seem to 
hold up when examined under field conditions. Al- 
though more study will be carried out on Ernodea 
in the Bahamas, under a tentative arrangement which 
seems reasonable at this stage, E. cokeri is considered 
to be a synonym of E. taylori, and E. nashii is merged 
with E. millspaughii. The latter name was chosen 
from the two originating at the same date of publi- 
cation because it is represented by the type specimen 
possessing both flowers and fruits. The type of E. 
taylori has no flowers at all; the type of E. nashii 
must have had flowers at the time Britton described 
the species, but they are missing from the type 





Rhodora [Vol. 76 

specimens now (types are at NY). The differences 
as observed in the type specimens of these micro- 
species seem inadequate for maintenance in this ex- 
ceedingly variable genus (also see Britton, 1908, for 
descriptions of “races”? within this complex). A key 
to identify the Bahamian species of Ernodea within 
my species concept at the moment follows: 

1. Calyx lobes nearly as long as the fruit, or 
longer .............. RR RRRRRRI 2. 

2. Corolla white to very pale pink .......... 
SENE Ernodea littoralis Sw. 

2. Corolla red ...... Ernodea angusta. Small. 
1. Calyx lobes less than half as long as the 
fruits 2.0... 0 eee 3. 

3. Leaves 6-8 mm. wide ................... 
MEIN Ernodea millspaughii Britton. 
3. Leaves 1-9 mm. wide ............... less 
ZEE Ernodea taylori Britton. 
Borreria saxicola > Borreria brittonii Standl. Un- 
like Ernodea, the eight species of Borreria in the 
Bahamas — or, at least most of them — seem to be 
good biological species with a high degree of ende- 
mism in the southern islands, especially Inagua and the 
Caicos group. Whereas actual treatment of this genus 
is deferred for the moment, one name change needs 
to be cited. Following Standley (1931), the name 
B. saxicola Britt. is replaced because it is a later 
Spermacoce — consult Bacigalupo (1972). 


Anguria pedata > Psiguria pedata (Jacq.) Howard. 
In his treatment of modern names for plants dis- 
cussed in Jacquin’s Selectarum, Howard (1978) 
found that the name Anguria Jacq. was a later 
homonym for Anguria Mil. The next available 
generic name is Psiguria; Howard made the neces- 
sary new combinations for this species, 










Bahama Flora — Gillis 121 
Lobelia — consult MeVaugh (1943). 
Scaevola plumierii > Scaevola plumieri (L.) Vahl. 

This orthographic change is made to conform to the 
present International Code. 


Xanthium chinense — Xanthium strumarium L. 
Xanthium strumarium is an earlier name than X. 
Ambrosia paniculata —^ Ambrosia artemisiifolia L. I 
have selected a lectotype of Ambrosia paniculata 
from material in the Michaux Herbarium at Paris. 
Furthermore, I have examined the lectotype of A. 
artemisiifolia which was selected by Payne (1970) 
at LINN, and concur with Payne that these plants 
are conspecific. The Linnaean name is obviously the 
older one and should be used. 
Iva — consult Jackson (1960). 
Ageratum latifolium —> Ageratum conyzoides subsp. 
latifolium (Cav.) M. F. Johnson. This name change 
has been made in accord with the treatment by John- 
son (1971). The typical subspecies has also been 
reported from the Bahamas, thus adding a taxon 
to the flora. 
Chrysopsis graminifolia > Heterotheca graminifolia 
(Michx.) Shinners. Shinners (1951), Wagenknecht 
(1960), and Harms (1964) have argued for the 
merger of Heterotheca and Chrysopsis. 
Aster bracei — Aster tenuifolius var. aphyllus Long. 
Long (1970a) has treated the Bahamian populations 
in connection with those of South Florida which he 
was studying. This taxon seems best treated as a 
variety of the more widespread A. tenuifolius. 
Leptilon linifolium — Conyza floribunda H.B.K. 

124 Rhodora [Vol. 76 


462 Juniperus lucayana — Juniperus bermudiana L. More 
work needs to be done on typification and on popula- 
tion studies in Juniperus to place the Bahamian plant 
properly. For the moment, it is treated in accord 
with Moore, 1966. 


463 A thorough monograph of Zamia is needed. In a 
manuscript by Chamberlain, left unpublished at his 
death, Z. lucayana has been referred to the Cuban 
Z. guttierezii Sauv. So little material of this plant, 
as well as of Z. angustifolia and Z. tenuis, is avail- 
able at this time, that it is not appropriate to make 
a nomenclatural judgment now. The recent redis- 
covery of the Long Island population of “Z. lucayana" 
by S. R. Hill (in press) and its subsequent cultiva- 
tion will help to make material available for further 


ACUNA, J. and J. T. Rorc. 1962, Psychotriae Cubenses Novae 
(Rubiaceae). Brittonia 14: 224-229. 

ADAMS, C. D. 1970. Miscellaneous additions and revisions to the 
flowering plants of Jamaica. Phytologia 20: 309-14. 

1971. Miscellaneous additions and revisions to the 

flowering plants of Jamaica. III. Phytologia 21: 405-10. 

1972. Flowering plants of Jamaica. University of 
the West Indies, Mona, Jamaica. 848 pp. 

ADAMS, WILLIAM P. 1957. A revision of the genus Ascyrum (Hy- 
pericaceae). Rhodora 59: 73-95. 

ADAMS, WILLIAM P. and NoRMAN K. B. RoBSON. 1961. A re-evalua- 
tion of the generic status of Ascyrum and Crookea (Gutttiferae). 
Rhodora 63: 10-16. 

ALAIN, HERMANO. 1962. Flora de Cuba, Vol. 5 — Rubiales - Valeri- 
anales - Cucurbitales - Campanulales - Asterales. Editorial Uni- 
versitaria, Universidad de Puerto Rico, Rio Piedras. 

ALLEN, CAROLINE K. 1966. Notes on Lauraceae of tropical America. 
I. The generic status of Nectandra, Ocotea, and Pleurothyrium. 
Phytologia 13: 221-27. 

1974] Bahama Flora — Gillis 125 

AMES, OAKES. 1910. A new Ponthieva from the Bahamas. Torreya 
10: 90-91. 

BACIGALUPO, NELINDA M. 1972. Observaciones sobre algunas es- 
pecies de los géneros Spermacoce L. y Spermacoceodes O.K. 
(Rubiaceae). Darwiniana 17: 341-57. 

BAEHNI, C. 1965. Mémoires sur les Sapotacées. III. Inventaire des 
genres. Boissiera 11: 1-262. 

BALDWIN, J. T., JR. 1938. Kalanchoé: the genus and its chromo- 
somes. Amer. J. Bot. 25: 572-79. 

Bates, DAvID M. 1965. Notes on the cultivated Malvaceae. I. Hi- 
biscus. Baileya 13: 56-130. 

BEAUVOIS, A. M. F. J. PaLisoT DE. 1812. Essai d'une nouvelle 
Agrostrographie. Paris. 

BECKNER, JOHN. 1970. Notes on Florida and West Indian orchids. 
Phytologia 20: 217. 

BELCHER, R. O. 1956. A revision of the genus Erechtites (Compos- 
itae) with inquiries into Senecio and Arrhenechthites. Ann. Mis- 
souri Bot. Gard. 43: 1-85. 

BoGLE, A. LINN. 1970. The genera of Molluginaceae and Aizoaceae 
in the southeastern United States, J. Arnold Arbor. 51: 431-62. 

BoRSSUM, WAALKES, J. VAN. 1966. Malesian  Malvaceae revised. 
Blumea 14: 1-213. 

BRASHIER, CLYDE K. 1966. A revision of Commelina (Plum.) L. 
in the U.S.A. Bull. Torrey Bot. Club 93: 1-19. 

BREMEKAMP, C. E. B. 1938. Notes on the Acanthaceae of Surinam. 
Rec. Trav. Bot. Neerl. 35: 130-170. + 4 pl. 

BRENAN, J. P. M. 1958. New and noteworthy Cassias from Tropical 
Africa. Kew Bull. 13: 231-252. 

BRITTON, N. L. 1908. The genus Ernodea Swartz: a study of 
species and races. Bull. Torrey Bot. Club 35: 203-208. 

1911. The botanical name of the wild sapodilla. 

Torreya 11: 128-29. 
1918. Flora of Bermuda. Charles Scribner's Sons. 

New York. 


Bahama flora. Published by the authors. Reprinted 1962, Hafner 

Publishing Co., New York. 

and JOSEPH NELSON RosE. 1928. Rosales - Mimo- 

saceae. North American Flora. Vol. 23, Pt. 1. 

1923-30. Scientific survey of Puerto Rico and the 
Virgin Islands. Botany of Puerto Rico and the Virgin Islands. 
Descriptive flora. Part 1 — New York Acad. Sci. 5 (1): 1-158; 
5 (2): 159-316; 5 (3): 317-474; 5 (4): 475-626; 6 (1): 1-158; 
6 (2): 159-316; 6 (3): 317-521; 6 (4): 523-663. 

BRIZICKY, GEORGE K. 1963. The genera of Sapindales in the south- 
eastern United States. J. Arnold Arbor. 44: 462-501. 

126 Rhodora [Vol. 76 

. 1964. The genera of Celastrales in the south- 

eastern United States. J. Arnold Arbor. 45: 206-34. 

1966. The genera of Sterculiaceae in the 

southeastern United States. J. Arnold Arbor. 47: 60-74. 

1968. Herissantia, Bogenhardia, and Gayo- 
ides (Malvaceae). J. Arnold Arbor. 49: 278-79. 

BuncH, DEREK. 1966. Two new species of Chamaesyce (Euphorbia- 
ceae), new combinations, and a key to the Caribbean members 
of the genus. Ann. Missouri Bot. Gard. 53: 90-9. 

CABRERA, ANGEL L. 1971. Revision del género Gochnatia (Com- 
positae). Revista Mus. de la Plata (N.S.— Botanica) 12: 1-160. 

CANDOLLE, A. P. DE. 1825. Prodromus. Vol. 2. Paris. 

CHANNELL, R. B. and C. E. Woop, Jr. 1959. The genera of the 
Primulales of the southeastern United States. J. Arnold Arbor. 
40: 268-88. 

CHASE, AGNES. 1929. The North American species of Paspalum. 
Contr. U.S. Natl. Herb. 28: 1-310 + xvii. 

1951. (revision of A. S. Hitchcock) Manual of the 
grasses of the Unied States, ed. 2. U.S.D.A. Misc. Publ. 200. 
1051 pp. 

CLAYTON, W. D. 1964. Studies in the Gramineae: vi. Sporoboleae. 
The Sporobolus indicus complex. Kew Bull. 19: 287-96. 

1968. The correct name of the common reed. 
Taxon 17: 168-69. 

CLEMENT, I. D. 1957. Studies in Sida (Malvaceae) 1. A review 
of the genus and monograph of the Section Malacroideae, Phy- 
salodes, Pseudomalvastrum, Incanifolia, Oligandrae, Pseudona- 
paea, Hookeria, and Steninda. Contr. Gray Herb. 180: 5-91. 

CoKER, W. C. 1944. The woody Smilaxes of the United States. J. 
Elisha Mitchell Soc. 60: 27-69. 

COLEMAN, JAMES R. 1966. A taxonomic revision of Section Ximene- 
sia of the genus Verbesina L. (Compositae). Am. Midl. Nat. 
76: 475-81. 

ConpiT, IRA J. 1969. Ficus, the exotic species. Division of Agri- 
cultural Sciences, University of California, Berkeley. 363 pp. 

CORCORAN, SISTER Mary Lucy. 1941. Revision of the Subgenus 
Pycreus in North and South America. Contr. Biol. Lab. Catholic 
Univ. Amer. 37: 1-68 + xvi. 

CORRELL, DONOVAN STEWART. 1950. Native orchids of North 
America, north of Mexico. Chronica Botanica Co., Waltham, 

CRISTOBAL, CARMEN L. 1960. Revision del género Ayenia (Ster- 
culiaceae). Opera Lilloana iv. Tucuman, Argentina, pp. 5-230. 

CRONQUIST, ARTHUR. 1943. "The separation of Erigeron from Conyza. 
Bull. Torrey Bot. Club 70: 629-632. 

1974] Bahama Flora — Gillis 127 

1944. Studies in the Simaroubaceae. IV. Ré- 
sumé of the American genera. Brittonia 5: 128-47. 

1945a. Studies in the Sapotaceae. I. The 
North American species of Chrysophyllum. Bull. Torrey Bot. 
Club 72: 192-205. 

. 1945b. Studies in the Sapotaceae III. Dipholis 
and Bumelia. J. Arnold Arbor. 26: 435-71. 

1945c. Studies in the Sapotaceae. IV. The 
North American species of Manilkara. Bull. Torrey Bot. Club 
72: 550-62. 

1946. Studies in the Sapotaceae. II. Survey of 

the North American genera. Lloydia 9: 241-92. 

1947. Notes on the Compositae of the north- 
eastern United States. V.  Astereae. Bull. Torrey Bot. Club 
74: 142-50. 

DANDY, J. E. 1967. Index of generic names of vascular plants, 
1753-1774. Reg. Veg. 51: 1-130. 

and A. W. EXELL. 1938. On the nomenclature of the 

species of Caesalpinia. J. Bot. 76: 175-80. 

and Geoffrey Tandy. 1939. On the identity of Syrin- 
godium Kütz. J. Bot. 77: 114-16. 

DARLINGTON, JOSEPHINE. 1934. A monograph of the genus Mentze- 
lia. Ann. Missouri Bot. Gard. 21: 103-226 + 6 pl. 

DELISLE, DONALD B. 1963. Taxonomy and distribution of the genus 
Cenchrus. Iowa State Univ. J. Sci. 37: 259-351. 

DEVALL, WILBUR B. 1941. The taxonomic status of Pinus caribaea 
Mor. Florida Acad. Sci. Proc. 5 (1940) : 121-32. 

DEWIT, H. C. D. 1955. A revision of the genus “Cassia” (Caesalp.) 
as occurring in Malaysia. Webbia 11: 197-292. 

. 1961. Typification and correct names of Acacia 
villosa Willd. and Leucaena glauca (L.) Bth. Taxon 10: 50-54. 

DEWOLF, GORDON P. 1960. Ficus. In: Flora of Panama. Ann. 
Missouri Bot. Gard. 47: 146-65. 

DRESSLER, ROBERT L. 1957. The genus Pedilanthus (Euphorbia- 
ceae). Contr. Gray Herb. 182: 3-188. 

. 1961. A reconsideration of Encyclia (Orchi- 
daceae). Brittonia 13: 253-66. 

DUNSTERVILLE, G. C. K. and LESLIE A. GARAY. 1965. Venezuelan 
orchids illustrated. Vol. 3. Andre Deutsch Ltd., London. 

EITEN, GEORGE 1963. Taxonomy and regional variation of Oxalis, 
section Corniculatae. Introduction, keys, and synopsis of the 
species. Amer. Midl, Nat. 69: 257-309. 

EPLING, CARL CLAWSON. 1925. Studies on South American Labia- 
tae. I. Synopsis of the genera Teucrium, Rosmarinus, Marru- 
bium, Prunella, Lamium, Leonurus, and Leonotis. Ann. Missouri 
Bot. Gard. 12: 107-129. 

128 Rhodora [Vol. 76 

1938-39. A revision of Salvia: Subgenus 
Calosphace. Fedde, Repert. Sp. Nov. 110: 1-380. 

ERNST, WALLACE R. 1963. The genera of Capparaceae and Morin- 
gaceae in the southeastern United States. J. Arnold Arbor. 44: 

FassETT, NORMAN C. 1955. Echinodorus in the American Tropics. 
Rhodora 57: 133-56; 174-88; 202-12. 

FERNALD, M. L. 1934. Reallignments in the genus Panicum. Rho- 
dora 36: 61-87. (also Contr. Gray Herb. 103). 

and K. M. WIEGAND. 1914. The genus Ruppia in 
eastern North America. Rhodora 16: 119-27. 

FRYXELL, PAUL S. 1971. A revision of Phymosia (Malvaceae). 
Madrono 31: 153-74. 

GALE, SHIRLEY. 1944. Rhynchospora, section Eurhynchospora, in 
Canada, the United States, and the West Indies. Rhodora 46: 
89-134; 159-97; 207-49; 255-78. (Contr. Gray Herb. 151). 

GAMERRO, JUAN CARLOS. 1968. Observaciones sobre la biología floral 
y morfología de la Potamogetonácea Ruppia cirrhosa (Petag.) 
Grande (R. Spiralis L. ex Dum.) Darwiniana 14: 575-608. 

GARAY, LESLIE A. and HERMAN R. SWEET. 1972. Notes on West 
Indian orchids III. J. Arnold Arbor. 53: 515-30. 

GILLIS, WILLIAM T. 1971. The systematics and ecology of poison- 
ivy and the poison-oaks (Toxicodendron, Anacardiaceae). Rho- 
dora 73: 72-237; 370-448; 465-540. 


1973. Additions to the Bahama flora since Britton and Mills- 

paugh — I. Rhodora 75: 411-425. 

and WILLIAM T. STEARN. in press. Typification 
and correct names of Leucaena and Lysiloma species in the Ba- 
hama flora. Taxon. 

GODFREY, R. K. 1952. Pluchea, section Stylimnus, in North Amer- 
ica, J. Elisha Mitchell Sci. Soc. 68: 238-71 + 23 pl. 

GOLDBERG, AARON. 1967. The genus Melochia L. (Sterculiaceae). 
Contr. U.S. Natl. Herb. 34: 191-363. 

GooDprNG, E. G. B., A. R. LovELESS, and G. R. PRocTOR. 1964. Flora 
of Barbados. Overseas Research Publication No. 7 (Ministry of 
Overseas Development). Her Majesty's Stationery Office. Lon- 
don. 486 pp. 

GouLD, FRANK W. 1967. The grass genus Andropogon in the 
United States. Brittonia 19: 70-76. 

- . 1968. Grass systematies. McGraw-Hill Co., New 
York. 382 pp. 

1969. The genus Schizachyrium in the Americas. 
XI. Int. Bot. Cong. Proc. p. 75. 

1974] Bahama Flora — Gillis 129 

GRAHAM, SHIRLEY A. and C. E. Woop, JR. 1965. The genera of 
Polygonaceae in the southeastern United States. J. Arnold Arbor. 
46: 91-121. 

GRENZENBACH, MYRTLE. 1926. A revision of the genus Bouchea 
(exclusive of the Chascanwm). Ann. Missouri Bot. Gard. 13: 
71-100 + 12 pl. 

GUNN, CHARLES R. 1972. Moonflowers, Ipomoea section Calonyc- 
tiom, in temperate North America. Brittonia 24: 150-68. 

HAGSTRÖM, J. O. 1916. Critical researches on the Potamogetons. 
Kungl. Sven. Vetens. Akad. Handl. Stockholms 55: 1-281. 

HAMET, R. 1907. Monographie du genre Kalanchoé. Bull. Herb. 
Boissier 7 (Sér. 2) : 869-900. 

HARMS, VERNON L. 1965. A second character distinguishing Het- 
erotheca s. str. from Chrysopsis (Compositae: Astereae). Rho- 
dora 67: 86-8. 

HARTOG, C. DEN. 1959. A key to the species of Halophila (Hydro- 
charitaceae), with descriptions of the American species. Acta 
Bot. Neerl. 8: 484-89. 

1964. An approach to the taxonomy of the sea- 

grass genus Halodule Endl. (Potamogetonaceae). Blumea 12: 


1970. The sea-grasses of the world. North-Holland 
Publishing Co., Amsterdam, 

HENRARD, J. TH. 1926-1933. A critical revision of the genus Aris- 
tida. Meded. Rijks. Herb. Leiden 54: 1-220 (1926); 54A: 221-464 
(1927); 54B: 465-701 (1928); 54C: 703-747 (1933). 

1929-1932. A monograph of the genus Aristida. 

Meded. Rijks. Herb. Leiden 58: 1-153 + Ix pl. (1929); 58A: 

157-325 + pl. Ixi-clix (1932). 

1950. Monograph of the genus Digitaria. Uni- 
versitaire Pers Leiden. 999 pp. 

HILL, STEVEN R. in press. Additions to the Bahama flora. Rhodora. 

HiTCHCOCK, A. S. 1936. Manual of the grasses of the West Indies. 
U.S.D.A. Misc. Publ. 243: 1-439. 

HiTrCHCOCK, C. LEO. 1932. A monographic study of the genus Ly- 
cium of the Western Hemisphere. Ann. Missouri Bot. Gard. 19: 
179-374 + 24 pl. 

. 1945. The Mexican, Central American, and 
West Indian Lepidia. Mandrofio 8: 118-43. 

HorLM, RicHARD W. 1950. The American species of Sarcostemma 
R. Br. (Asclepiadaceae). Ann. Missouri Bot. Gard. 37: 447-560. 

Horvat, SISTER MARY LiGOURI. 1941. A revision of the subgenus 
Mariscus found in the United States. Contr. Biol. Lab. Catholic 
Univ. Amer. 33: 1-147 + ix. 

130 Rhodora [Vol. 76 

Horcukiss, NEIL and HERBERT L. DoziER. 1949. Taxonomy and 
distribution of North American cat-tails. Amer. Mid. Nat. 41: 

HOWARD, RICHARD A. 1957. Studies in the genus Coccoloba. IV. 
The species from Puerto Rico and the Virgin Islands and from 
the Bahama Islands. J. Arnold Arbor. 38: 211-42. 

1962. Notes on Buxus in the Lesser Antilles 

and on Mathou's overlooked publication. J. Arnold Arbor. 44: 


1973. The Enumeratio and Selectarum of 

Nicolaus von Jacquin. J. Arnold Arbor. 54: 435-470. 

and HENRY F. DUNBAR. 1964. Additions to 

the flora of Inagua, the Bahamas. Rhodora 66: 6-15. 

and D. A. POWELL. 1968. Clerodendrum phil- 
ippinum Schauer replaces “Clerodendrum fragrans.” Taxon 17: 

HUTCHINSON, J. B. 1943. The cottons of Jamaica. Trop. Agr. 20: 

INGRAM, JoHN. 1967. A revisional study of Argythamnia, subgenus 
Argythamnia (Euphorbiaceae). Gentes Herb. 10: 1-38. 

IsELY, DUANE. 1970. Legumes of the United States. II. Desman- 
thus and Neptunia. Iowa State J. Sci. 44: 495-511. 

1972. Legumes of the United States. VI. Calliandra, 
Pithecellobium, Prosopis. Madrofio 21: 273-98. 

JACKSON, Curtis R. 1949. A key to the genus Scleria Berg in 
South Florida. Quart. J. Florida Acad. Sci. 12: 220-22. 

Jackson, R. C. 1960. A revision of the genus Iva L. Univ. Kansas 
Sci. Bull. 41: 793-876. 

JERVIS, Roy NEWELL. 1954. A summary of the genus Gochnatia 
including a revision of the West Indian species which comprise 
the Section Amastraphioides. Unpublished Ph.D. Dissertation. 
University of Michigan. Ann Arbor, Michigan, 228 pp. 

JoHNSON, Mites F. 1971. A monograph of the genus Ageratum L. 
(Compositae — Eupatorieae). Ann. Missouri Bot. Gard. 58: 6-88. 

JOHNSTON, IvAN M. 1949a. Studies in the Boraginaceae xvii. A. 
Cordia section Varronia in Mexico and Central America. J. 
Arnold Arbor. 30: 85-110. 

1949b. The botany of San José Island (Gulf 
of Panama). Sargentia 8: 1-306. 

JOHNSTON, MARSHALL. 1962. The North American mesquites, Pro- 
sopis section Algarobia (Leguminosae). Brittonia 14: 72-90. 
1964. The fourteen species of Ziziphus in- 
cluding Sarcomphalus (Rhamnaceae) indigenous to the West 

Indies. Amer. J. Bot. 51: 1113-15. 

1971. Revision of Colubrina (Rhamnaceae). 

Brittonia 23: 2-53. 

1974] Bahama Flora — Gillis 131 

KEARNEY, THOMAS H. 1954a. A tentative key to the North Amer- 
ican species of Pavonia Cav. Leafl. West. Bot. 7: 122-30. 

1954b. A tentative key to the North Amer- 

ican species of Sida L. Leafl. West. Bot. 7: 138-50. 

1954c. Notes on Malvaceae V. Leafl. West. 

Bot. 7: 118-21. 

KILLIP, ELLSWORTH P. 1938. The American species of Passiflora- 
ceae. Field Mus. Nat. Hist. Publ. Bot. Ser. 19 (1), Publ. 407. 

KOBUSKI, CLARENCE EMMEREN. 1926. A revision of the genus Priva. 
Ann. Missouri Bot. Gard. 13: 1-344- 5 pl. 

Kopp, LUCILLE E. 1966. A taxonomic revision of the genus Persea 
in the Western Hemisphere (Perseae — Lauraceae). Mem. N.Y. 
Bot. Gard. 14: 1-120. 

KRAL, RoBERT. 1971. A treatment of Abildgaardia, Bulbostylis, and 
Fimbristylis (Cyperaceae) for North America. Sida 4: 57-227. 

KUKENTHAL, G. 1949. Vorarbeiten zu einer Monographie der Rhyn- 
chosporideae. Bot. Jahrb. 74: 375-509. 

KunTH, C. S. 1824. Synopsis Plantarum aequinctialium orbis novi 
(Voyage de Humboldt et Bonpland), Vol. 3. Paris. 

LAKELA, OLGA. 1963. Centrosema floridanum (Britton) Lakela, 
comb. nov. (Leguminosae). Sida 1: 182. 

Lay, Ko Ko. 1950. The American species of Triumfetta L. Ann. 
Missouri Bot. Gard. 37: 315-95. 

LEENHOUTS, P. W. 1967. A conspectus of the genus Allophylus 
(Sapindaceae). Blumea 15: 301-358. 

LEFOR, MICHAEL WILLIAM. 1968. A revision of the genus Roche- 
fortia Sw. (Boraginaceae). Unpublished M.S. thesis, University 
of Connecticut. Storrs. 

LEGRAND, C. D. 1952. Revisando tipos de Portulaca. Communic. 
Bot. Mus. Hist. Nat. Montevideo 2 (24): 1-10 + 2 pl. 

LEON, HERMANO. 1946. Flora de Cuba. Vol. 1 Gimnospermas y 
Monocotiledoneas. Contr. Ocas. Mus. Hist. Nat. Colegio de la 
Salle. No. 8 Habana. 

LEON, HERMANO and HERMANO ALAIN. 1951-57. Flora de Cuba. 
Contribucion No. 10 — Dicotiledoneas: Casuarinaceas a Meliaceas. 
Vol. 2. Contr. Ocas. Mus. Hist. Nat. Colegio de la Salle. Habana. 
Contribucion No. 13 — Dicotiledoneas: Malpighiaceae a Myrta- 
ceae. Contribucion No. 16 — Dicotiledoneas: Malastomaeaceae a 
Plantaginaceae. Vol. 4. 

LEWis, WALTER H. 1961. Merger of the North American Houstonia 
and Oldenlandia under Hedyotis. Rhodora 63: 216-23. 

LINNAEUS, C. 1769. Plantarum Jamaicensium Pugillus. Disserta- 
tion 102. 

1775. Plantae Surinamensis. Jac. Alm. Dissertation 


132 Rhodora [Vol. 76 

LITTLE, E. L., JR. 1947. The name of the wild dilly of Florida. 
Rhodora 49: 289-93. 

1968. Transfers to Guapira from  Torrubia 
(Nyctaginaceae). Phytologia 17: 367-71. 

Lowc, RoBERT W. 1970a. Additions and nomenclatural changes in 
the flora of southern Florida — I. Rhodora 72: 17-46. 

1970b. The genera of Acanthaceae in the south- 
eastern United States. J. Arnold Arbor. 51: 257-309. 

LÜCKHOFF, H. A. 1964. The natural distribution, growth, and bo- 
tanical variation of Pinus caribaea and its cultivation in South 
Africa. Ann. Univ. Stellenbosch, Ser. A: 39: 1-60. 

LUER, CARLYLE A. 1972. The native orchids of Florida. New York 
Botanical Garden. New York. 293 pp. 

MCCANN, CHARLES. 1945. Notes on the genus Ruppiaceae. J. Bom- 
bay Nat. Hist. Soc. 45: 396-402. 

McGivney, SISTER M. VINCENT DE PAUL. 1938. Revision of the 
subgenus Eucyperus found in the United States. Contr. Biol. 
Lab. Catholic Univ. Amer. 26: 1-74 + xvii. 

MCLAUGHLIN, ANNE DesmMonp. 1944. The genus Cyperus in the 
West Indies. Catholic Univ. Amer. Biol. Studies No. 5 pp 1-108 
+ viii. 

McVaucu, Rocers. 1943. Campanulales — Campanulaceae (Lobeli- 
oideae). North American Flora 32A, Pt. 1: 1-124. 

1963. Tropical American Myrtaceae. II. Notes 

on generic concepts and descriptions of previously unrecognized 

species. Fieldiana 29: 393-532. 

1973. Notes on West Indian Myrtaceae. J. 
Arnold Arbor. 54: 309-14. 

MACBRIDE, J. FRANCIS. 1916. Further notes on the Boraginaceae. 
Contr. Gray Herb. 49: 16-22. 

1930. Other Peruvian plants, chiefly new 
species. Publ. Field Mus. Nat. Hist, Bot. Ser. (Publ. 278) 8: 

MERRILL, E. D. 1910. An enumeration of Philippine Leguminosae 
with keys to genera and species. (Concluded). Philip. J. Sci. 
(Botany) 5: 95-136. 

1936. On the application of the binomial Amaran- 

thus viridis Linnaeus. Amer. J. Bot. 23: 609-12. 

and Surv-YiNa Hu. 1949. Work and publications of 

Henry Muhlenberg, with special attention to unrecorded or in- 

correctly recorded binomials. Bartonia 25: 1-66. 

and F. P. METCALF. 1942. Hedyotis Linnaeus versus 

Oldenlandia Linnaeus and the status of Hedyotis lancea Thun- 

berg in relation to H. consangwinea Hance. J. Arnold Arbor. 23: 


1974] Bahama Flora — Gillis 133 

MOHLENBROCK, ROBERT H. 1957. A revision of the genus Stylosan- 
thes. Ann. Missouri Bot. Gard. 44: 299-355. 

MoLDENKE, HaRoLD N. 1936. A monograph of the genus Callicarpa 
as it occurs in America and in cultivation. ii. Fedde, Repert. Sp. 
Nov. 40: 38-131. 

1937. A monograph of the genus Petitia. 

Fedde, Repert. Sp. Nov. 42: 229-51. 

1958a. Materials toward a monograph of the 

genus Citharexylum II. Phytologia 6: 262-320. 

1958b. Materials toward a monograph of the 

genus Citharexylum III. Phytologia 6: 322-368. 

1958c. Materials toward a monograph of the 

genus Citharexylum IV. Phytologia 6: 383-432. 

1965. Materials toward a monograph of the 

genus Lippia. I. Phytologia 12: 6-71. 

1971. A fifth summary of the Verbenaceae, 
Avicenniaceae, Stilbaceae, Dicrastylidaceae, Symphoremaceae, 
Nyctanthaceae, and Eriocaulaceae of the world as to valid taxa, 
geographic distribution, and synonymy. Privately published, 
Wayne, NJ. 

Moore, HanoLD E., Jr. 1963. An annotated checklist of cultivated 
palms. Principes 7: 119-84. 

1966. Nomenclatural notes on the cultivated 

conifers. Baileya 14: 1-11. 

and WILLIAM T. STEARN. 1967. The identity 
of Achras zapota L. and the names for the sapodilla and the 
sapote. Taxon 16: 382-95. 

MoonE, R. J. and C. FRANKTON. 1969. Cytotaxonomy of some Cir- 
sium. species of the eastern United States, with a key to eastern 
species. Canad. J. Bot. 47: 1257-75. 

NICKERSON, Norton H. and JoskPH W. Tripp. 1973. Floral di- 
morphism in Rachicallis americana (Jacq.) Hitchc. (Saltwater- 
bush). Rhodora 75: 111-13. 

NIKLES, DoNALD GARTH. 1966. Comparative variability and rela- 
tionship of Caribbean Pine (Pinus caribaea Mor.) and slash pine 
(Pinus elliottii Engelm.). Unpublished Ph.D. dissertation. North 
Carolina State University. Raleigh, N.C. 

O'DoNELL, C. A. 1941. Revision de las especies americanas de 
Merremia (Convolvulaceae). Lilloa 6: 467-554. 

OGDEN, E. C. 1943. The broad-leaved species of Potamogeton of 
North America north of Mexico. Rhodora 45: 57-105; 119-63; 
171-214 + pl. 146-48. 

O'NEILL, HucH T. 1939. The status and distribution of Cyperus 
distinctus Steud. Rhodora 41: 131-33. 

134 Rhodora [Vol. 76 

1940. The sedges of the Yucatan Peninsula. xix. 
in Botany of the Maya area. Carnegie Inst. Washington Publ. 
522: 248-322. 

OosTSTROOM, S. J. VAN. 1934. A monograph of the genus Evol- 
vulus. Meded. Bot. Mus. Herb. Rijks. Univ. Utrecht No. 14: 

1940. The Convolvulaceae of Malaysia III. 

The genus Ipomoea. Blumea 3: 481-582. 

in collaboration with R. D. HooGLanp. 1953. 
Convolvulaceae in: Flora Malesiana I: 4: 388-512. 

ORNDUFF, RoBERT. 1969. Neotropical Nymphoides (Menyanthaceae): 
Meso-American and West Indian species. Brittonia 21: 346-52. 

Pacut, JiRí. 1952. Synopsis of the genus Catalpa (Bignoniaceae). 
III. Candollea 13: 241-85. 

PARKS, JAMES C. 1973. A revision of North American and Carib- 
bean Melanthera (Compositae). Rhodora 75: 169-210. 

PAYNE, WILLARD W. 1964. A re-evaluation of the genus Ambrosia 
(Compositae). J. Arnold Arbor. 45: 401-38. 

PENNELL, FRANCIS W. 1917. Notes on plants of the southern United 
States — III. Bull. Torrey Bot. Club 44: 337-62. 

Perry, JAMES D. 1971. Biosystematic studies in the North Amer- 
ican genus Sabatia (Gentianaceae). Rhodora 73: 309-69. 

PFEIFER, Howarp W. 1966. Revision of the North and Central 
American hexandrous species of Aristolochia (Aristolochiaceae). 
Ann. Missouri Bot. Gard. 53: 115-96. 

1970. Taxonomic revision of pentandrous 
species of Aristolochia. University of Connecticut Publication 
Series. Storrs, Connecticut. 

PuiLcox, D. 1965. Revision of the New World species of Buchnera 
L. (Serophulariaceae). Kew Bull. 18: 275-315. 

PoRTER, DUNCAN M. 1972. The genera of Zygophyllaceae in the 
southeastern United States. J. Arnold Arbor. 53: 531-52. 

POWELL, A. MICHAEL. 1965. Taxonomy of Tridax (Compositae). 
Brittonia 17: 47-96. 

PRANCE, GHILLEAN T. 1972. Chrysobalanaceae. Flora Neotropica 
Monograph No. 9. Hafner Publishing Co. New York. 

PRESL, C. B. 1835. Reliquiae Haenkeanae, Vol. 2. Prague. 

PURSEGLOVE, J. W. 1968. Tropical crops.  Dicotyledons. 2 vol. 
Longmans, London. 719 pp. 

RAO, ARAGULA SATHYANARAYANA. 1956. A revision of Rauvolfia, 
with particular reference to the American species. Ann, Missouri 
Bot. Gard. 43: 253-354. 

RAVEN, PETER H. 1962. New combinations in Ludwigia. Kew Bull. 
15: 476. 

1974] Bahama Flora — Gillis 135 

1968. The Old World species of Ludwigia (In- 
cluding Jussiaea) with a synopsis of the genus (Onagraceae). 
Reinwardtia 6: 327-427. 

Reap, RoBERT W. 1968. A study of Pseudophoenix (Palmae). 
Gentes Herb. 10: 169-213. 

REED, CLYDE F. 1968. The nomenclature and synonymy of the 
Amaranthaceae, Chenopodiaceae, and Nyctaginaceae of the flora 
of Texas. Phytologia 18: 1-44. 

REESE, G. 1962. Zur intragenerischen Taxonomie der Gattung Rup- 
pia L. Zeit. für Bot. 50: 237-64. 

ROBERTSON, KENNETH R. 1971. The Malpighiaceae in the south- 
eastern United States. J. Arnold Arbor. 52: 101-12. 

Rosson, N. 1965. New and little known species from the Flora 
Zambesiaca area. XVI. Taxonomic and nomenclatural notes on 
Celastraceae. Bol. Soc. Broter. 39: 5-55. 

RoE, KEITH E. 1968. Solanum verbascifolium L., misidentification 
and misapplication. Taxon 17: 176-78. 

Rogers, C. MARVIN. 1963. Yellow flowered species of Linum in 
eastern North America. Brittonia 15: 97-122. 

1968. The Linwm bahamense complex. Rhodora 

70: 439-41. 

Rogers, Davip J. 1963. Studies on Manihot esculenta Crantz and 
related species. Bull. Torrey Bot. Club 90: 43-54. 

RoMINGER, JAMES M. 1962. Taxonomy of Setaria in North Amer- 
ica. Illinois Biol. Monogr, 29: 1-132. 

ROTHMALER, WERNER. 1944. Nomina generica neglecta 1753-1763. 
Fedde, Repert. Sp. Nov. 53: 1-37. 

Rupp, VELVA E. 1967. Oxyrhynchus and Monoplegma (Legumino- 
sae). Phytologia 15: 289-94. 

1968. Leguminosae of Mexico — Faboideae. T 

Sophoreae and Podalyrieae. Rhodora 70: 492-532. 

1969. A synopsis of the genus Piscidia (Legumino- 
sae). Phytologia 18: 473-99. 

St. JoHN, HanoLp. 1970. Classification and distribution of the 
Ipomoea pes-caprae group (Convolvulaceae). Bot. Jahrb. 89: 

SAUER, JONATHAN. 1964. Revision of Canavalia. Brittonia 16: 


1972. Revision of Stenotaphrum (Gramineae: 
Paniceae) with attention to its historical geography. Brittonia 
24: 202-22. 

SAVAGE, SPENCER. 1945. A catalogue of the Linnaean herbarium. 
Linnaean Society of London. London. 225 pp. 

ScHERY, ROBERT WALTER. 1942. Monograph of Malvaviscus. Ann. 
Missouri Bot. Gard. 29: 183-244 + 17 pl. 

136 Rhodora [Vol. 76 

ScHINDLER, A. K. 1926. Die Desmodiinen in der botanischen Liter- 
atur bis Linné. Fedde, Repert. Sp. Nov. 23: 71-127. 

ScuuLz, Orro E. 1909. Citharexylum Linn. Sym. Antill. 6: 56-69. 

ScuwaNiTZ G. 1967. Untersuchungen zur Postmeiotischen Mikro- 
sporogenese. 1. Morphogenese des Ruppia-pollens. Pollen et 
Spores 9: 9-48. 

SEALY, J. ROBERT. 1954. Review of the genus Hymenocallis. Kew 
Bull. 1954: 201-40. 

SETCHELL, W. A. 1946. The genus Ruppia. Proc. Calif. Acad. Sci. 
4a. S 25: 469-72. 

SHERFF, EARL EDWARD. 1937. The genus Bidens. Field Mus. Nat. 
Hist. Bot. ser. 16: 1-709. 

1945. Some additions to the genus Dodon- 

aea L. (Sapindaceae). Amer. J. Bot. 32: 202-14. 

1947. Further studies in the genus Dodon- 
aea L. (family Sapindaceae). Publ. Field Mus. Bot. 23: 269-317. 

SHINNERS, L. H. 1951. The North Texas species of Heterotheca, 
including Chrysopsis. Field and Lab. 19: 66-71. 

1957. Synopsis of the genus Eustoma (Gentian- 

aceae) Southwestern Nat. 2: 38-43. 

1962. Micromeria brownei and its allies (Labi- 
atae). Sida 1: 94-97. 

SLEE, M. U. 1970. Crossability values within the slash-Caribbean 
Pinus species complex. Euphytica 19: 184-89. 

SPRAGUE, T. A. 1932. Laugeria “Vahl” = Terebraria Kuntze. Kew 
Bull. 1926: 349. 

STAFLEU, FRANS A. 1967. Taxonomic Literature. Regnum Vegeta- 
bile 52. 556 pp. Utrecht. 

STAFLEU, FRANS et al. 1972. International Code of Botanical No- 
menclature. Reg. Veg. 82: 1-426. 

STEARN, WILLIAM T. 1957. The boat-lily (Rhoeo spathacea). Bai- 
leya 5: 195-98. 

1958. A key to West Indian mangroves. Kew 

Bull. 1958: 33-37. 

1964. Catharanthus roseus, the correct name 
of the Madagascar periwinkle. Lloydia 27: 196-200. 

1966. Botanical Latin. Thomas Nelson and 
Sons, Ltd. London. 566 pp. 

1968. Jamaican and other species of Bumelia 
(Sapotaceae). J. Arnold Arbor. 49: 280-89. 

1969. A synopsis of Jamaican Myrsinaceae. 
Bull. Brit. Mus. Nat. Hist. Bot. 4: 143-78 -- 8 pl. 

1971. A survey of the tropical genera Oplonia 
and Psilanthele (Acanthaceae). Bull. Brit. Mus. Nat. Hist. Bot. 
4: 259-323. 

1974] Bahama Flora — Gillis 137 

1972. Typification of Evolvulus nummularius, 
E. convolvuloides, and E. alsinoides (Convolvulaceae). Taxon 21: 

STUCKEY, RONALD L. 1972. Taxonomy and distribution of the genus 
Rorippa (Cruciferae) in North America. Sida 4: 279-430. 

TAYLOR, PETER. 1967. Lentibulariaceae. Mem. N.Y. Bot. Gard. 17: 

THARP, B. C. and MARSHALL C. JOHNSTON. 1961. Recharacteriza- 
tion of Dichondra (Convolvulaceae) and a revision of the North 
American species. Brittonia 13: 346-60. 

TRELEASE, W. 1918. Agave in the West Indies. Mem. Natl. Acad. 
Sci. 11: 1-55. 

1916. The genus Phoradendron: a monographie re- 
vision. Univ. Illinois Press. Urbana. 

VERDCOURT, B. 1963. Convolvulaceae. in: Flora of Tropical East 
Africa, C. E. Hubbard and E. Milne-Redhead. eds.; Crown Agents 
for Overseas Governments and Administrations. 

WAGENKNECHT, BURDETT L. 1960. Revision of Heterotheca, section 
Heterotheca (Compositae). Rhodora 62: 61-76; 99-107. 

WATERFALL, U. T. 1967. Physalis in Mexico, Central America, and 
the West Indies. Rhodora 69: 82-102; 203-40; 319-29. 

WEBSTER, GRADY L. 1956-58. A monographie study of the West 
Indian species of Phyllanthus. J. Arnold Arbor. 37: 91-122; 217- 
68; 340-59; pls. 1-14; 38: 51-80; 170-98; 295-373; pls. 15-23; 39: 
49-100; 111-212; pls. 24-32. 

1967. The genera of Euphorbiaceae in the 

southeastern United States. J. Arnold Arbor. 48: 303-61; 363-430. 

1970. A revision of Phyllanthus (Euphorbi- 
aceae) in the continental United States. Brittonia 22: 44-76. 

WHEELER, Louis C. 1938. The names of three species of Brassica. 
Rhodora 40: 306-309. 

WIJNANDS, D. O. 1973. Typification and nomenclature of two spe- 
cies of Sansevieria (Agavaceae). Taxon 22: 109-114. 

WILBUR, ROBERT L. 1955. A revision of the North American genus 
Sabatia (Gentianaceae). Rhodora 57: 1-33; 43-71; 78-104. 
WILSON, KENNETH A. 1960. The genera of Convolvulaceae in the 
southeastern United States. J. Arnold Arbor. 41: 298-317. 
WINDLER, D. R. 1966. A revision of the genus Neptunia (Legum- 

inosae). Aust. J. Bot. 14: 379-420. 

Woop, CARROLL E., JR. 1958. The genera of the woody Ranales in 
the southeastern United States. J. Arnold Arbor. 39: 296-346. 

1959. The genera of the Nymphaeaceae and 

Ceratophyllaceae in the southeastern United States. J. Arnold 

Arbor. 40: 94-112. 

138 Rhodora [Vol. 76 

and R. B. CHANNELL. 1960. The genera of 
the Ebenales in the southeastern United States. J. Arnold Arbor. 
41: 1-35. 

Woopson, ROBERT E., JR. 1936. Studies in the Apocynaceae, IV. 
the American genera of Echitoideae. Ann. Missouri Bot. Gard. 
23: 169-438 + 7 pl. 

1937. New or otherwise noteworthy Apo- 
cynaceae of tropical America, V. Ann. Missouri Bot. Gard. 24: 

1938a. Studies in the Apocynaceae. VII. 
An evaluation of the genera Plumeria L. and Himatanthus 
Willd. Ann. Missouri Bot. Gard. 25: 189-224. 

1938b. (Asclepiadales) Apocynaceae. 
North Amer. Flora 29: 103-192. 

YAKOVLEV, G. P. 1967a. Systematical and geographical studies of 
the genus Sophora L. and allied genera. Prob. Pharmacog. 21: 
42-62. (In Russian). 

1967b. Some characteristics of structure of seeds 
of Sophora L. and allied genera in connection with its systematics 
and phylogeny. Repr. Proc. Leningrad Chem.-Pharm. Inst. 21: 
90-98. (In Russian). 

YATES, HARRIS O. 1966. Revision of grasses traditionally referred 
to Uniola I. Uniola and Leptochloópsis. Southwestern Nat. 11: 

YUNCKER, TRUMAN GEORGE. 1932. The genus Cuscuta. Mem. Tor- 
rey Bot. Club 18: 113-331. 



To those who had always considered Witmer Stone as 
an ornithologist — and one of the country's greatest — the 
appearance in 1911 of his “Plants of Southern New Jersey" 
came as a distinct surprise. Had they known more about 
the man they would have realized that he was one of the 
last of the all-around naturalists, interested in mammals, 
insects and plants as well as birds. 

The recent facsimile reprint of Stone's book, by the 
Quarterman Publications, Inc., of Boston, makes available 
once more a work which for more than sixty years has been 
the standard reference for everyone who is interested in 
the flora of the Pine Barrens, as well as the other plant 
provinces of southern New Jersey. 

Originally published as Part II of the Annual Report 
of the New Jersey State Museum, this important volume 
has for many years been out of print and has, indeed, be- 
come something of a collector's item. 

Although one can not be other than grateful that this 
flora is once again in circulation, it is to be regretted that 
the Foreword was not written by a professional botanist, 
preferably one who was intimately acquainted with Witmer 

Since this reprint is a facsimile, every word, every punc- 
tuation mark and every misprint appears exactly as it did 
in the original. There are numerous discrepancies between 
its nomenclature and that in use today. It would have been 
helpful to the reader if Elizabeth M. Woodford, who wrote 
the Foreword, had explained the reasons for this. 

Partly because he was a zoologist and partly because of 
his friendship with Dr. N. L. Britton of the New York 
Botanical Garden, Stone's treatment of plant names follows 
the now-abandoned American Code of Botanical] Nomen- 

'Stone, Witmer. The Plants of Southern New Jersey. Reprint. 
1973. Quarterman Publications, Inc., Boston, Massachusetts. 828 pp. 
Illust. $25.00. 


140 Rhodora [Vol. 76 

clature. This code, like that of the zoologists, permits the 
use of tautonyms or duplicate binomials. Hence we find 
in Stone such combinations as Blephariglottis blephari- 
glottis, Hepatica hepatica, Linaria linaria and others. 

The American Code never embraced the principle of the 
conservation of generic names and Stone therefore con- 
sistently employed what he considered to be the earliest 
generic epithets. As a consequence we have scores of 
generic names which are unfamiliar to most present-day 
taxonomists. A few examples will suffice: Spathyema for 
Symplocarpus, Juncoides for Luzula, Abama for Narthe- 
cium, Vagnera for Smilacina, Gyrostachys for Spiranthes, 
Hicoria for Carya, Ammodenia for Arenaria (peploides), 
Meibomia for Desmodium, etc. 

Fortunately most of the presently accepted names are 
cited in synonymy (although not always with the correct 
authority), but it seems a pity that a republication of this 
work should not have suggested some revision and up- 
dating of its nomenclature. It might also have corrected 
such misprints as “Sandy Hood” for “Sandy Hook" (page 
432) and “dandelon” for “dandelion” (page 817). 

Mrs. Woodford states that Stone’s collection of 12,000 
specimens went to the herbarium of the Academy of Natu- 
ral Sciences of Philadelphia. Such is not the case. Realizing 
that many of his duplicates were already in the Academy 
and that he had frequently been in the field with members 
of the Philadelphia Botanical Club whose specimens had 
likewise been deposited there, I suggested to Dr. Stone that 
he leave his herbarium to the University of Pennsylvania. 
He readily agreed, and in the fall of 1942 the University 
received his collection of more than 14,000 sheets, rich not 
only in plants from southern New Jersey, but also in ma- 
terial from northern Pennsylvania. It was my pleasant 
task to supervise the incorporation of this collection into 
the herbarium of the University. 

Witmer Stone once told me that his manuscript for the 
New Jersey Flora had been prepared in a little more than 
four months. Doubtless he had made some preliminary 

1974] Book Review — Fogg 141 

efforts to put together his observations accumulated over 
many years of field work, but even so this was a prodigious 
accomplishment. He added that the Director of the New 
Jersey Museum had approached him in desperation to give 
them something for their Annual Report and urged him 
to prepare a paper on the Pine Barrens of New Jersey. 
Stone's reply was that this would be like treating one 
vegetational province in a vacuum and that he preferred 
to deal with the entire flora of the lower three-fifths of 
the state, at the same time emphasizing the geographic 
origins of the various elements represented therein. This 
he did superbly well, in a manner which certainly would 
have elicited the approbation of Asa Gray. 

Mrs. Woodford gives a picture of Witmer Stone as a 
warm, friendly human being with a wide breadth of in- 
terests and a well developed sense of humor. With this i 
can heartily agree. It was my good fortune for many 
years to spend my summers at Cape May, where Dr. Stone 
also had a cottage, and in many walks with him I came 
to appreciate his encyclopedic knowledge and great capacity 
for friendship. 

MERION, PA. 19066 


A new series of monographie studies, Ginkgoana,' was 
inaugurated early in 1973 with the publication of Number 1, 
“The Asiatic species of Desmodium and its allied genera 
(Leguminosae)," by Hiroyoshi Ohashi of the Department 
of Botany, University of Tokyo. 

The first study in the series is truly a book, printed in a 
clear and pleasing type face, and very well edited. The 
text is written in English. This study is a portion of the 
thesis presented by Mr. Ohashi for the degree of Doctor of 
Science. It is an inclusive treatment with clearly stated 

Since the genus Desmodium (in the broad sense) is one 
in which I have long had a particular interest and since I 
know the Asiatic species less well than the American and 
African, I am pleased to see this detailed piece of work. 
Of special importance are the pollen studies and the meticu- 
lous investigations of floral parts, seeds, and seedlings. 
Dr. Ohashi confirms the conclusions of Ambrose? for Des- 
modium (i.e. the species of Desmodium ser. Americana have 
hypogaeous germination, while those of other species ob- 
served have an epigaeous type of germination), and pre- 
sents new information for additional species. 

The chapter considering observations on morphological 
characters demonstrates well the care and scope of Dr. 
Ohashi's observations on habit, leaves, inflorescences, bracts, 
bracteoles, androecia or stamens, pollen grains, fruits 
[*^pods"], seeds, seedlings, and chromosome numbers. He 
has illustrated with clarity the basic structures and prob- 
able evolutionary progression in the development of the 

'Ginkgoana, Contributions to the Flora of Asia and the Pacific 
Region. No. 1, The Asiatic Species of Desmodium and its Allied 
Genera (Leguminosae), by Hiroyoshi Ohashi, 318pp., 86 figures, 76 
plates [18.1 X 25.6 em.]. February 15, 1973. Academia Scientific 
Book Ine., Tokyo, Japan. 

"The Michigan Botanist 6: 97-99. 1967. 


1974] Book Review — Schubert 143 

inflorescence for the group (Desmodium sens. lat.) and also 
of the bracts and bracteoles. In both cases the tendencies 
indicated could be compared profitably with parallel or 
similar progressions in the New World and African groups 
of related genera. The resulting comparisons, particularly 
with New World genera and species, should be of much 
interest because of the numerous groups of closely related 
species which radiate from the centers of diversity in Mex- 
ico and Brazil. 

The taxonomie treatment has well constructed keys, de- 
tailed descriptions, ample citations of specimens, exceed- 
ingly good illustrations of floral parts and of fruits, and 
distribution maps. In general, I think Dr. Ohashi has rather 
over-classified the group he has studied so carefully, follow- 
ing Schindler and Hutchinson in large part. However, he 
has documented his data and stated his reasons so carefully 
that his occasionally debatable conclusions do not detract 
from the usefulness of the work. The treatment of Des- 
modium subgenus Sagotia is perhaps the one which troubles 
me most. Dr. Ohashi says “The subgenus Sagotia is one of 
the most polymorphic and widespread groups of the genus 
Desmodium. It is characterized by having mostly the ter- 
minal racemes composed of 2(-3)-flowered fascicles which 
are subtended usually only by the primary bracts (i.e. lack- 
ing the secondary bracts), ebracteolate flowers, diadelphous 
androecia and rather small-sized articles of the fruits.” He 
includes, however, species with secondary bracts, among 
them Desmodium velutinum, D. gangeticum, and D. dichoto- 
mum; some with bracteoles, such as D. caudatum and D. 
benthamii; some with flowers borne singly; and some with 
loment articles of considerable size. 

It seems to me that the only reason the genus Sagotia 
(sensu Ohashi) is so polymorphic is that Dr. Ohashi has 
assigned to it very unlike elements which do not have a 
close natural relationship. His arrangement may seem 
feasible when studying only Asian species, but on a world 
wide basis I think it would become wholly unworkable and 
its defects would be much magnified. In some infrageneric 

144 Rhodora [Vol. 76 

groups in Asia as for most of the genus Desmodiwm in 
Africa there are many unrelated, distinctive species in con- 
trast to the situation in North and South America where 
there are many small to large groups of closely related 
species. This makes for a very different aspect of a pre- 
sumed phylogeny, of proposed evolutionary series, and for 
an understanding of interspecific relationships in Asia and 
Africa on the one hand and in America on the other, even 
though there are some species which bridge the gap by 
natural or artificial means. 

The only serious omissions in this careful study are in 
the areas of ecology and phytogeography, discussion of the 
former being almost totally lacking and of the latter only 
minimal. This may indicate a lack of field experience which 
can be remedied later by an active field program. Field 
work might also give Dr. Ohashi a broader view of the taxa 
with which he is working and a somewhat different ap- 
proach to their interrelationships. 

In concluding this review I would stress the meticulous 
care of the observations Dr. Ohashi has made and the great 
gap he has filled in our knowledge of one large complex in 
the family Leguminosae. 




One of the most important tasks of botanists specializing 
in taxonomy and geobotany is the compilation of flora 
manuals for their colleagues in general and for the much 
more numerous amateur botanists in particular. In this 
way, the author himself gets a supplementary survey over 
the plants of the region in which he works and gathers 
experience that enables him to solve problems of unusual 
importance. At the same time he gets a rare opportunity 
to stimulate interest in his subject and to gather new prose- 
lytes among the younger generation to carry on in his foot- 
steps. Only a few botanists have the encyclopedic knowl- 
edge and ability to write floras, and still fewer want to take 
the risk of being unfairly criticized for this least gratifying 
of scientific occupations, because in no other field is it easier 
to be criticized for opinions and judgements. This is the 
perennial trouble for compilers of floras and also the main 
cause of the scarcity of good manuals. If the author also 
ventures to make deviations from the accepted delimita- 
tions, descriptions and names of the plants included, he 
runs the additional risk of condemnation by his more con- 
ventional colleagues, because scientists are no less con- 
servative than are other citizens. Ordinary manuals are 
always valuable, but unconventional floras ought to be 
doubly welcome, because nothing else can stimulate future 
research to a higher degree. 

Although Switzerland is one of the smaller European 
countries, it has fostered astonishingly many of the botani- 
cal giants. Several of its botanists certainly have been con- 
ventional and followed the trend of their times, whereas 
others have dared to have new ideas and have looked at the 

'Hans Ernst Hess, Elias Landolt, Rosemarie Hirzel: Flora der 
Schweiz und angrenzender Gebiete. Three volumes, 29 X 23 cm., 
2690 pages, with 9 colored plates and about 3450 drawings. Birk- 
háuser Verlag, Basel & Stuttgart, 1967, 1970, 1972. Price 450 
Schweizer francs. 


146 Rhodora [Vol. 76 

plants of their country from new points of view. These 
latter we have to thank for the unusually large number of 
presently well-known species that were first described from 
Switzerland. Although many of the great Swiss botanists 
of the past are now always mentioned with reverence, there 
were times when their contemporaries called them names 
and were critical about their scientific abilities. Such is the 
fate of those who are born too early or try to be prophets 
in their own land, but time is their confederate. 

It would be difficult to mention all the many flora man- 
uals, which have been published in Switzerland in the past, 
and the list of those which have reached fame far outside 
the country because of scientific quality is far from being 
short. One such flora was Albrecht von Haller’s great 
*Historia stirpium indigenerorum Helvetiae inchoata", 
which was published in 1768 in three folio volumes. That 
work has stood in a class by itself for two centuries as the 
most monumental botanical manual ever written in the field 
of descriptive floristies. It still remains important for its 
clear descriptions and other information, but the conserva- 
tism in nomenclature and the reluctance to accept what 
would be the future in that field has detracted from the 
nomenclatural importance of the work though it always will 
be a text of literary superiority rarely achieved in scientific 

Although many good Swiss floras have been published 
since Haller's work, two centuries had to pass before a text 
of similar magnitude and quality again was compiled for 
this flora of more than 3500 species. This is the new “Flora 
der Schweiz," by H. E. Hess, E. Landolt and R. Hirzel, the 
three large volumes of which were published in 1967, 1970 
and 1972. It is a work of extraordinary quality and quan- 
tity, based on fifteen years of intensive work by the two 
first authors and on equally intensive work of drawing the 
excellent pictures of more than 3450 of the species by the 
third author. 

The virtues of this manual are too many to be even sim- 
ply listed, and readers with different inclinations will al- 

1974] Book Review — Lóve 147 

ways find some matters of special interest. Considerable 
information on the methods and philosophy adopted in the 
compilation of this great work is given in an introductory 
chapter, which then is followed by almost thirty pages on 
the history and origin of the flora and its distribution, and 
more than forty pages on its ecology. Specialists are apt 
to find omissions in these chapters, and some may perhaps 
join issue with the authors as to some of the explanations 
given. That is to be expected in any work of this magnitude 
and concentration. But this reviewer, who believes he has 
gathered considerable knowledge about this flora through 
visits and reading numerous good books and reports during 
the past three decades, finds these chapters to be highly 
educational and of great interest from more points of view 
than he cares to mention. Numerous maps in these intro- 
ductory chapters ought to make it easy for those unfamiliar 
with the geography of the land to follow the detailed ac- 

The manual itself is arranged according to the Engler 
system as are most modern European floras. A general key 
to the higher groups leads to keys to the families of the 
major groups where these commence, and keys to genera 
and species are then given in appropriate places in the 
text. Families and genera are described in detail, followed 
under the latter by information about the size of the genus 
and its distribution, and about cytological information avail- 
able. After the generic key there are detailed reports of 
possible hybridization observed, frequently with explana- 
tions of the morphological, geographical, and cytological 
characteristics of the plants. 

Most species are represented by a good drawing, fre- 
quently with additional details. Their Latin names are 
followed by synonyms, if any, and German names, and a 
very detailed description is given of the morphological 
characteristics. For every species there is also information 
on chromosome number and other cytological peculiarities, 
and the authors have done an admirable work in looking up 
the original papers in order to be able to exactly inform the 

148 Rhodora [Vol. 76 

reader about the geographical origin of the material. This 
is the only place where such information has been compiled, 
and it is admirably corect, although some few mistakes have 
not been avoided. Then there are reports on the ecological 
occurrence of each species in Switzerland, and on the gen- 
eral distribution of the taxon elsewhere in the world. Fre- 
quently, the authors then add taxonomical and geographical 
remarks on the variation of the species within and outside 
their country, especially when they feel that it is to be 
regarded as a critical taxon in need of more close investiga- 
tion, with references to the most recent literature in every 
special case. 

It is easy to find points of disagreement in a work of this 
magnitude, although this reviewer is of the opinion that the 
authors have sueceeded unusually well in avoiding harsh 
judgements. They have made an effort to accept a modern 
species concept although sometimes this may not have suc- 
ceeded well enough, and the descriptions of the taxa are 
clear and moderately concise. It is a pity that they have 
found it impossible to mention subspecific categories, but 
since Switzerland abounds in such variations which fre- 
quently may be of doubtful value, it is understandable that 
the authors avoided spending perhaps twice as much time in 
evaluating these. The only serious remark to be made seems 
to be that the authors adopted, for present-day European 
botanists, an unusually conservative attitude towards the 
nomenclature, so that sometimes their eccentricity may look 
disturbing. Nevertheless, this is understandable, and the 
explanation of this given in the last volume is not unrea- 
sonable although I believe that a more liberal attitude would 
have made the work better and also more useful for a longer 
period of time. 

After the description of the last species in the third vol- 
ume, there are more than 90 pages of corrections and addi- 
tions, half a page with an epilogue, where some criticism of 
plant sociology and nomenclature by reviewers of the first 
two volumes are met, and four pages on which the termi- 
nology used is explained. There are good indexes of Latin 

1974] Book Review — Lóve 149 

and German names, and these are preceded by sixty pages 
of a very important bibliography. 

The three volumes are printed in the unusual size 29 X 
23 em, with considerable empty space on every page that is 
partially used for the large drawings. The heavy volumes 
are strongly and tastefully bound in green linen, and they 
are certainly reasonably priced for their size and quality. 
Their weight would make it difficult to carry them into 
the field, but the authors are compiling a smaller manual 
with the keys only and new drawings of selected species 
that will be a handy companion on excursions in the Alps 
and elsewhere. 

The “Flora der Schweiz” is an unusually thorough and 
scholarly work which will long be the standard reference 
for this part of Europe. It is also an invaluable source of 
information for botanists from other parts of the continent, 
and also for specialists in other fields, especially that of 
cytotaxonomy. The text and pictures are so detailed and 
exact that the book can be highly recommended to those 
foreign botanists and institutions which need good and 
reliable information on European plants which may have 
been introduced overseas, because no other flora comple- 
ments the comprehensive Flora Europaea more excellently 
for southern and central European plant species. 



Manuscripts must be double-spaced or preferably triple- 
spaced (not on corrasable bond), and a list of legends for 
figures and maps provided on a separate page. Footnotes 
should be used sparingly, as they are usually not necessary. 
Do not indicate the style of type through the use of capitals 
or underscoring, partieularly in the citations of specimens, 
except that the names of species and genera may be under- 
lined to indicate italies in discussions. Specimen citations 
should be selected critically especially for common species 
of broad distribution. Systematic revisions and similar 
papers should be prepared in the format of “The System- 
aties and Ecology of Poison-Ivy and the Poison-Oaks," 
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu- 
larly with reference to the indentation of keys and syno- 
nyms. Papers of a floristic nature should follow, as far as 
possible, the format of ‘Contribution to the Fungus Flora 
of Northeastern North America. V.," H. E. Bigelow & M. E. 
Barr, Rhodora 71: 177-203. 1969. For bibliographic cita- 
tions, a recommended list of standard journal abbreviations 
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. 
Club 85: 277-300. 1958. 

Volume 76, No. 805, including pages 1-150, was issued March 30, 1974. 


CONTENTS: — continued 

Studies on the Bryophytes of Southern Manitoba III. 
Collections from Grand Beach Provincial Park 

Paul W. Stringer and Muriel H. L. Stringer .................... 27 
Some New or Noteworthy Vascular Plant Records from 

Northwestern Saskatchewan 

Vernon L. Harms and John H. Hudson ...............5.4....« 39 
Parasitic Witchweed: Striga asiatica Versus S. lutea 


F. N. Hepper eee eene ennt ent tna th tta anne tnntn seen tine th atas tna ta nan 45 
Cacalia rugelia: A New Combination for a North American 


T. M. Barkley and Arthur Cronquist ............. eee 48 
Typification and Application of the Names Scirpus ameri- 

canus Pers. S. olneyi Gray, and S. pungens Vahl. 

Alfred E. Schuyler ............ eee eee eee eee eene te nent tn nennen nnn 51 
The Correct Authority for Cardamine clematitis 


y JU C209 207,177 NERS EE EATE AA IE E E T 53 
A New Form of Digitaria sanguinalis 

John E. Ebinger .rsccssosssssssessssssssecccserscsssnesensseterevssesacccssssssescescnns 57 
The Growth of some New England Perennial Seaweeds 

Narman EEO ER aai iori d diassec itaq E a 59 
Chromosome Numbers in Kosteletzkya Presl. (Malvaceae) 

Orland J. Blanchard, Jv. sscsssccersssescersrsssccccoveccessonconecssenseseceees 64 
Name Changes for Seed Plants in the Bahama Flora 

William T. Gillis eaea aa eaan A 67 
Book Review 

JORN EOI: IT: aeien oicseeri tis reei  E 139 
Book Review 

Bornio G uSOhWDOT 5 un udis aszsessensoes iic iaa neem: 142 

Book Review 




Conducted and published for the Club, by 


œ Associate Editors 


Vol. 76 June, 1974 No. 806 


A Systematic Treatment of the Native Unifoliolate Crotalarias 
of North America (Leguminosae) 

Donald R. Wiéndler ....cccccrccccscescsssssercosscessconessscssecceccssscessccosasences 151 
The Flora of Mount Monadnock, New Hampshire 
Er o e E A E T E E E T 205 

Taxonomy of Perityle Section Perityle 
NH MB AR eia A E A E ER A E A E 229 

Carex exilis, Geocaulon lividum and Other Plants of Interest 
in Pittsburg, New Hampshire 
BEEN TONTO n A n a N eiii 307 

(Continued on Inside Back Cover) 

The Nem England Botanical Club, Ine. 
Botanical Museum, Oxford St., Cambridge, Mass. 02138 

RHODORA. — A quarterly journal of botany, devoted primarily to the 
flora of North America and floristically related areas. Price $10.00 
per year, net, postpaid, in funds payable at par in the United States 
currency at Boston. Some back volumes, and single copies are 
available. For information and prices write RHODORA at address 
given below. 

Scientific papers and notes, relating directly or indirectly to the 
plants of North America, will be considered by the editorial com- 
mittee for publication. Articles concerned with systematic botany 
and cytotaxonomy in their broader implications are equally accept- 
able. All manuscripts should be double-spaced throughout. Please 
conform to the style of recent issues of the journal. Illustrations 
can be used only if the cost of engraver's blocks is met through the 
author or his institution. Forms may be closed five weeks in advance 
of publication. Extracted reprints, if ordered in advance, will be 
furnished at cost. 

Address manuscripts and proofs to A. Linn Bogle, 

Dept. of Botany and Plant Pathology, 
University of New Hampshire, 
Durham, New Hampshire 03824 
Subscriptions and orders for back issues (making all remittances 
payable to RHODORA) should be sent to RHODORA, Botanical 

Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive 

the next number of RHODORA, changes of address must be received 

prior to the first day of March, June, September or December. 
Second Class Postage Paid at Boston, Mass. 




Vol. 76 June, 1974 No. 806 



Crotalaria is a large genus of tropical, subtropical, and 
temperate plants comprised of about 550 species, over 400 
of which occur in Africa. In 1939 Senn reported 31 species 
for North America with the majority growing in Mexico 
and the West Indies; only 21 of these species were con- 
sidered to be native. Eight of the 21 species, including the 
Linnean C. sagittalis, form a group that is so distinctive 
that Rafinesque (1837) separated it into the subgenus 
Iocaulon. The Iocaulon? Crotalarias are annual or perennial 
plants characterized by simple leaves; decurrent stipules, 
when present; terminal or leaf-opposed inflorescences ; and 
glabrous, inflated fruits that become black at maturity. 
Senn (1939) indicated that this is the most difficult group 
of North American Crotalarias in which to delimit species 
and suggested the need for further work. 

'A portion of a dissertation submitted as partial fulfillment of the 
requirements for the Doctor of Philosophy Degree in the Department 
of Botany, University of North Carolina at Chapel Hill. 

"Contribution number 7 from Towson State College Herbarium. Cost 
of Publication of illustrations assisted by the Towson State College 
Faculty Research Fund. 

"The name Iocaulon will be used as an adjective throughout this 
paper to refer to the unifoliolate Crotalarias related to C. sagittalis. 


152 Rhodora [Vol. 76 

The history of the classification of Crotalaria has been 
reviewed by Baker (1914), Senn (1939), and Polhill 
(1968). While Baker treated the African species of Crota- 
laria primarily, he also reviewed the history of the classi- 
fication of the genus and discussed its separation from the 
closely related genus Lotononis. Senn reviewed rather 
briefly the nomenclatural history of Crotalaria in North 
America, and discussed the placement of the North Amer- 
ican species into sections described by Baker. Senn's treat- 
ment includes a key to the species, nomenclature and 
synonomy, Latin descriptions for newly described species, 
lists of specimens examined, and a few comments on each 
of the taxa. Most species lack descriptions and all lack 
illustrations, which makes it difficult to identify a plant if 
the key only narrows the choice to a group of species, 

Polhill (1968), while dealing primarily with the African 
species of Crotalaria, presented a discussion of the history 
of generic subdivisions, discussed typification of the genus, 
and proposed a new arrangement of species into sections 
based on combinations of flower characters. The characters 
used were: (1) type and position of standard appendages, 
(2) twisting of the keel, (3) lobing of the calyx, (4) calyx- 
keel length ratio, (5) pubescence of the style, (6) style 
shape, and (7) receptacle size. Polhill placed C. sagittalis 
and its relatives in the section Calycinae Wight & Arnott. 
All the characters that Polhill used for this placement were 
found to be consistently present in the Iocaulon Crotalarias 


In addition to the personal collections made in the United 
States and Mexico, specimens were borrowed from twenty- 
one herbaria. The names and locations of the herbaria 
from which specimens were borrowed appear below, ac- 
companied by the abbreviation of each as listed in Index 
Herbariorum, except in one case, marked by an asterisk, 
where no listing could be found for the herbarium and an 
arbitrary abbreviation is used. 

1974] Crotalarias — Windler 153 

A— Arnold Arboretum, Cambridge, Massachusetts. 
ALU — University of Alabama, University, Alabama. AUA — 
Auburn University, Auburn, Alabama. GA — University of 
Georgia, Athens, Georgia. GH — Gray Herbarium, Cam-. 
bridge, Massachusetts. LAF — University of Southwestern 
Louisiana, Lafayette, Louisiana. MICH — University of 
Michigan, Ann Arbor, Michigan. *MISS — University of 
Mississippi, University, Mississippi. NCU — University of 
North Carolina, Chapel Hill, North Carolina. NY — The 
New York Botanical Garden, New York, New York. SIU — 
Southern Illinois University, Carbondale, Illinois. TEX — 
University of Texas, Austin, Texas. UC — University of 
California, Berkeley, California. us — United States Na- 
tional Museum, Washington, D.C. usF— University of 
South Florida, Tampa, Florida. wis — University of Wis- 
consin, Madison, Wisconsin, 

BM — British Museum of Natural History, London, Eng- 
land. G-— Conservatoire et Jardin Botanique, Geneva, 
Switzerland. K-— Royal Botanic Gardens, Kew, Great 
Britain. MEXU — Universidad Nacional de México, Mexico 
City, Mexico. P — Museum Nacional d'Histoire Naturelle, 
Paris, France. 

Crotalaria plants from more than 60 locations in the 
United States and Mexico were grown from seed at the 
North Carolina Botanical Garden, Chapel Hill, during the 
summer of 1968. Data from the garden plantings were 
used in evaluating the status of taxa. Another paper de- 
tailing the results of the field and garden studies is published 
elsewhere (Windler, 1973). 


CROTALARIA (Dillenius ex Linnaeus, Hort. Cliff. p. 218, 
1731, pro parte) L., Sp. Pl. p. 714 pro maxima parte. Type 
species: Crotalaria lotifolia L. (see Polhill, 1968 pp. 174-5). 

Erect, ascending, spreading, decumbent, or procumbent 

154 Rhodora [Vol. 76 

terrestrial herbs or shrubs. Root a taproot with slender 
branches, nodules frequently present. Stems single in the 
erect species to many in the decumbent and procumbent 
species, the epidermis green, frequently becoming purple, 
with an indument varying from nearly glabrous to serice- 
ous, of spreading or appressed, simple trichomes, Stipules 
absent or varying from a pair of subulate lobes projecting 
from near the leaf base to inverted-sagittate, decurrent, 
foliaceous appendages which wing the stem for a portion 
of the subtending internode, a full internode, or for several 
of the subtending internodes, the lobes spreading, incurved 
toward the stem or parallel to the stem. Leaves alternate, 
unifoliolate, undivided, entire, ciliate, membranaceous to 
fleshy, broadly ovate to lanceolate or linear, 0.9-10.2 cm. 
long, 0.3-4.6 cm. wide, the adaxial surface dark green, gla- 
brous, or with spreading or appressed, simple trichomes, 
the abaxial surface usually lighter green than the adaxial 
and with spreading or appressed simple trichomes; petioles 
0.5-3 mm. long. 

Inflorescences loose racemes, peduncles borne terminally, 
or opposite leaves and 0.0-3 cm. below the node. Bracts 
lanceolate, elliptic, subulate, or linear, to 14 mm. long, one 
at the base of each flower, frequently persistent after flower 
or fruit abscission. Flowers papilionaceous. Calyces bilabi- 
ate, deeply cleft, the upper two lobes free and broader than 
the lower three, the lower usually remaining coherent, the 
calyx bearing two lateral bracteoles, one on either side. 
Standards (vexillum, banner) orbicular, obovate, or ellip- 
tic-oblong, the apex rounded or retuse, 4 mm. shorter to 6 
mm. longer than the upper calyx lobes, yellow, the adaxial 
surface frequently becoming red-tinged, lined or unlined, 
nearly glabrous with a few trichomes along the main vein 
on the adaxial surface, with lamelliform appendages at the 
base of the abaxial surface of the blade. Wings (alae) ob- 
long, attached by a stalk continuous with the lower margin 
(cleaver-shaped), the blades usually puckered between some 
of the veins, yellow. Keel (carina) petals twisted at the 
tip, white or greenish-white, lanate-ciliate. Stamens 10, 

1974] Crotalarias — Windler 155 

monadelphous, the staminal tube split on the upper side, 
anthers dimorphie, with basifixed long anthers alternating 
with medifixed short anthers, the short anthers positioned 
above the elongate at anthesis. Styles geniculate at the 
base, bearing a spiral line of trichomes transversing the 
length and culminating in a tuft of trichomes on the stigma. 

Fruit an inflated, sub-cylindric, short-stipitate legume, 
glabrous inside and out, becoming black at maturity. Seeds 
7-46 per pod, oblique-cordiform, smooth. 

The systematic treatment presented here covers twelve 
native North American unifoliolate species of Crotalaria 
section Calycinae. The descriptions are based on materials 
collected during the study and on numerous herbarium 
specimens. Measurements of stipule width are taken at 
the base of the sinus between the two lobes and include the 
width of both at that point. Length measurements of the 
calyx and standard are made from the base of the torus to 
the tip of the respective structure in the undissected flower. 
Illustrations were prepared by Mrs. Jerry Meyer. 

Since data on cross-fertility between the numerous 
populations sampled is not available, taxa have been dis- 
tinguished primarily on patterns of morphological discon- 
tinuity. It is assumed that these discontinuities also reflect 
breeding discontinuities, especially in areas where species 
are sympatric. In mountainous areas, however, many pop- 
ulations are well isolated, frequently leading to morpholog- 
ical discontinuity between populations. In these instances 
greater emphasis has been placed on similarities in inter- 
preting relationships. 

Diagnostic characters that have been useful in the group 
are plant habit; general size; position of inflorescence; 
presence, absence, size, and shape of the stipules; length 
and position of trichomes; and flower size. Most of the 
diagnostic characters are present on the plants during the 
entire growing season. 

The synonomy includes only those names that were pos- 
sible to check either against types or clear descriptions. A 


Rhodora [Vol. 76 

special effort is made to point out all places where the 
present treatment differs from that of Senn's. l 

A set of duplicate specimens from the author’s field col- 
lections has been deposited in the herbarium of the Univer- 
sity of North Carolina at Chapel Hill (NCU). 



A. Inflorescence terminal, or terminal and also leaf- 

opposed. ...........leeee RR heh B. 
B. Plants with decurrent stipules. (West Indies, Cen- 

tral and South America). ........... 1. C. pilosa. 
B. Plants without decurrent stipules. ............ C. 

C. Shrubs and subshrubs with diffuse habit. .. D. 

D. Peduncles 9 cm. or more long; calyx 15-20 
mm. long. (Mexico, Central and South 
America). .......... esses 2. C, nitens. 

D. Peduncles 6 cm. or less long; calyx 10-11 
mm. long. (Northwestern Mexico). ..... 

3. C. brevipedunculata. 

C. Herb with decumbent or ascending habit. 

(Western Mexico). ........ 4. C. polyphylla. 

A. Inflorescence not terminal, always leaf-opposed. ..... 
eee e hehe ehe E. 
E. Plants erect or strongly ascending. ........... F. 
F. Stem pubescence spreading; adaxial leaf sur- 
faces regularly pubescent. ............... G. 

G. Stipules with lobes recurved. (West Indies 

and South America). .... 5. C. stipularia. 

G. Stipules with lobes spreading, (Through- 

out). eee eee ee eee 6. C. sagittalis. 

F. Stem pubescence appressed, adaxial leaf sur- 

faces glabrous or with appressed trichomes. 

H. Upper leaf surface glabrous. ......... I. 
I. Plants perennial; stipules usually con- 
spicuous; peduncles 7.5-23 cm. long. 

1974] Crotalarias — Windler 157 

(Southern United States). .......... 

NEP EL o M NC 7. C. purshii. 

I. Plants annual; stipules absent or incon- 
spicuous; peduncles 2.5-8 cm. long. 

(Mexico and Central America). ...... 

"T c IEEE 8. C. quercetorum. 

H. Upper leaf surface bearing trichomes. .. J. 

J. Peduncles 0.5-2 cm. long, usually as 

thick as the stem; stipules present; 
trichomes dense, obscuring the stem. 

HMETICO)S S.V VI 9. C. mexicana. 

J. Peduncles 2.5 cm. long or longer, thinner 

than the stem; stipules absent; tri- 

chomes not obscuring the stem (except 
occasionally near the top). (Southern 

United States). .. 10. C. rotundifolia. 

E. Plants decumbent, procumbent or spreading. .. K. 
K. Mature plants decumbent or procumbent from 
a thick taproot, stems usually less than 6 dm. 
long; trichomes on stem 0.3-3.2 mm. long. 
(Mexico and Southeastern United States). .. 
eer er ere o o 10. C. rotundifolia. 

K. Mature plants diffuse or spreading, stems usu- 
ally more than 6 dm. long; trichomes on stem 

usually less than 1 mm. long. (Mexico). .. L. 
L. Calyx 13-20 mm. long; stipules 0.5-2.7 cm. 
broad. (Mexico). .. 11. C. bupleurifolia. 
L. Calyx 7.5-12 mm. long; stipules narrow or 
absent. (Mexico). .. 12. C. nayaritensis. 

1. Crotalaria pilosa Miller, Garden Dict. No. 2. 1768, non 
C. pilosa Roxb. ex Mart. Denkschr. Konigl. Akad. Wiss. 
München 6: 156, 1920, non C. pilosa Thunb. Prod. Pl. 
Capensium 125. 

Crotalaria pterocaula Desv. J. Bot. (Desvaux) 3: 76. 
1814. Type: not located. 

Crotalaria genistella HBK. Nov. Gen. et. Sp. Pl. 6: 398. 
1824. Type: Humboldt 2050 (P, mc Microfiche observed). 

158 Rhodora [Vol. 76 

ac a 

Figure 1. Crotalaria pilosa. 
A. Habit and leaf variation 
B. Stem 


Crotalarias — Windler 


Su2QWw '*7) 'suo2vM 

‘DUNAS '? pue soyd +4 ‘syion16 ‘IBA 

‘IVA SUazU 7140]0304/) jo uonnqujsuiq ‘T dey 

.. 06 001 On 
| Blueindijs'9 o | | 
eSo|Id') e | | 
Ssijioe48 uea suajiu *) X | 
su3]iu ') 9 
j | |o 
hí a ee 
we $ o | 
— 1 ia | 
‘ NE. ® go | 
f MSN l 
a) f ? X | 
Qo. Tr, | 
q | 
4. | 
$ | ` | 
1 | re 
Mw | | 
| | | 
| | 
| | $ | 
I| 1 Ñ 
\ Ü e 
| | x 
| | | 
\ ? | n 
\ A | | ct 
pe b MED 
oL 08 06 ~ 001 E oll 

160 Rhodora [Vol. 76 

Crotalaria pilosa Miller var. skutchii Senn, Rhodora 41: 
331. 1939. Type: Skutch 3071 (GH). 

Annual erect herb with a taproot to 0.4 cm. thick. Stem 
single, 2.8-9.5 dm. long, 1.3-4 mm. thick, the longest inter- 
node 2.8-8.2 cm. long; vestiture of appressed or loosely ap- 
pressed trichomes 0.5-1.5 mm. long. Stipules present, de- 
current more than the length of the internode, 0.3-0.8 cm. 
wide, the width constant, the lobes 2.1-9.7 mm. long, usually 
not spreading. Leaves elliptic, lanceolate, narrowly lanceo- 
late, or linear, 3.3-8.6 cm. long, 3.5-25 mm. wide, base 
rounded to cuneate, the apex acute; vestiture of both sur- 
faces of appressed trichomes 0.4-2 mm. long; petioles 1-1.3 
mm. long. Inflorescences terminal or leaf-opposed and ter- 
minal, peduncles 4.5-12 cm. long. Bracts sessile, linear, or 
subulate, 3.1-4 mm. long, 0.3-0.5 mm. wide; pedicels 3-4.8 
mm. long. Calyces 9-11.5 mm. long, the tube 2.5-3.1 mm. 
long, with appressed, loosely appressed, or somewhat 
spreading trichomes 0.5-1 mm. long; bracteoles elliptic, 
narrowly elliptic, or subulate, 2.5-4 mm. long, 0.4-0.5 mm. 
wide. Corollas yellow, the standards 9-12.8 mm. long, 
equaling or to 1.7 mm. longer than the upper calyx lobes; 
elongate anthers 1.5-2 mm. long, short anthers 0.3-0.5 mm. 
long ; style 5.5-5.8 mm. long. Fruits 2.5-3.6 cm. long, breadth 
not known; seeds tan, brown, or red-brown, 15-23 per pod, 
2.9.5 mm. long. Chromosome number: n—16. Flowering 
time: July-March. Habitat: sandy and gravelly soils in 
fields and open pine woods, elevation sea level to 2,400 feet. 
Range: Costa Rica, Cuba, Jamaica, Panama, (Also widely 
distributed in South America.) Figure 1, Map 1. 

Type: “La Vera Cruz in New Spain" (Mexico, Cuba or 
Jamaica?), Houstoun (BM; photo., NY!). 

In spite of the fact that Miller, in his original descrip- 
tion, cited Crotalaria pilosa as “growing naturally at La 
Vera Cruz in New Spain", no specimens have been observed 
from Veracruz, Mexico. The citation appears to be in 
error. The probable true type locality was either in Cuba 
or Jamaica. Senn, in his treatment of C. pilosa, recognized 
the species as being composed of three varieties. Senn's 

1974] Crotalarias — Windler 161 

variety skutchii is here treated as being a part of variety 
pilosa; his variety robusta is transferred to C. bupleurifolia 
and treated as a variety of that species. No field collec- 
tions were made of C. pilosa during this study and no seed 
was obtained for garden plantings. 

Crotalaria pilosa is characterized by its erect habit, ter- 
minal or leaf-opposed and terminal inflorescences, and 
stipules that wing the stem for more than one node below 
the node of origin (Figure 1). Occasional specimens have 
spreading lobes at the summit of the stipules. Such speci- 
mens observed are Britton, Britton, & Shafer 734 (NY) 
and Bro. Leon & Fr. M. Rosca 8837 (NY), both from Cuba. 
Crotalaria pilosa is alone among the species covered in this 
treatment in possessing both terminal inflorescences and 

San Ramon y Otevas, Brenes 11332 (NY). CUBA. Isle of Pines: 
pinelands, Britton, Britton, and Wilson 14615 (NY, US); PINAR DEL 
RIO: near Rio de las Vueltas, N of S. Diego de los Banos, Bro. Leon 
guana and Guerra, Howard and Howard 9924 (cH). JAMAICA. 
Upper Clarendon, James Hill Savannah, Harris 12844 (GH, NY, US). 
PANAMA. Ancon Hill, Canal Zone, open hillside, Killip 12075 (us). 
TRINIDAD. Without date or specific locality, de Suber 369 (P). 

2. Crotalaria nitens HBK. Nov. Gen. et Sp. 6: 399. 1824. 
Crotalaria bracteata Schlecht. et Cham. Linnaea 5: 575. 
1830, non Roxburgh, Hort. Bengal 54. 1814. (Type: 
Schiede 597, Hacienda de la Laguna, (B; photo., F!). 
Crotalaria schiedeana Steudel, Nomen. Botanicus. 1: 445. 
1840. Type: same as for C. bracteata for which this was 
a nomen nov. 

Shrub with unknown root type. Stems 5-15 dm. long, 2-4 
mm. thick, the longest internode 1-5.5 em. long; vestiture 
of appressed or loosely appressed trichomes, 0.5-1 mm. long. 
Stipules absent. Leaves elliptic, lanceolate, oblong, or 
oblanceolate, 2.5-7.5 (10) em. long, 3.5-18.5 mm. wide, base 
rounded to cuneate, the apex rounded, acuminate, or acute; 
vestiture of both surfaces of appresed or loosely appressed 

162 Rhodora [Vol. 76 

trichomes, 0.3-1 mm. long; petioles 1.3-3 mm. long. Inflor- 
escenses terminal, peduncles 6-51 cm. long. Bracts petiolate, 
lanceolate, the petioles 0.3-1 mm. long, the blades 4-14 mm. 
long, 0.6-3 mm. wide; pedicels 4.1-7 mm. long. Calyces 
15-18 (20) mm. long, the tubes 3.3-5 mm. long, bearing 
appressed or loosely appressed trichomes 0.3-1 mm. long; 
bracteoles lanceolate, elliptic-lanceolate or linear, 4.5-9 mm. 
long, 0.5-1.4 mm. wide. Corollas yellow, the standards 
13.5-19 (22) mm. long, equaling to 2.5 mm. longer than the 
upper calyx lobes; elongate anthers 2.2-3.6 mm. long, short 
anthers 0.6-1.2 mm. long; styles 8.5-11.5 mm. long. Fruits 
2.5-3.2 em. long, breadth not known; seeds grey-brown, 
brown, or red-brown, 26-40 per pod, 1.8-2.8 mm. long. 
Chromosome number: not known. Flowering time: Sep- 
tember-December. Habitat: bare soil, dry open mountain- 
sides and pinelands, elevation 4,000-7,200 feet. Range: 
Guatemala; Mexico: Chiapas, Jalisco, Oaxaca, Veracruz 
(Also widely distributed in South America). 

Type: Mariquita (Colombia), Humboldt (P, nc Micro- 
fiche observed). 

A photograph of the type of Crotalaria nitens was ob- 
served in the Ipc microfiche photographs of the Humboldt, 
Bonpland, and Kunth herbarium (P). It bears only the 
name C. nitida and the location Mariquita which is also 
mentioned in the location sites for C. nitens. It compares 
favorably with the photograph examined by Senn (1939) 
of the “co-type” in the Willdenow Herbarium. Senn’s appli- 
cation of the name is followed in this treatment. 

Crotalaria nitens is characterized by its shrubby habit, 
lack of stipules, terminal inflorescences, and large flowers. 
Of the North American species, it most resembles C. brevi- 
pedunculata but differs from it in having a larger flower 
size and longer peduncles. 

Two varieties are recognized: 
Peduncles stout, 1-2 mm. thick; bracts 7.5-14 mm. long, 
1.2-3 mm. wide. .......... 2a. C. nitens var. nitens. 
Peduncles slender, 0.5-0.6 mm. thick, bracts 4-5 mm. long, 
0.6-0.8 mm. wide. ........ 2b. C. nitens var. gracilis. 

1974] Crotalarias — Windler 163 


(hi i | 
y Wt 



a Z 

Figure 2. Crotalaria nitens var. nitens. 
A. Habit and leaf variation 

B. Stem 
Crotalaria nitens var. gracilis. 
C. Habit 

D. Stem 

164 Rhodora [Vol. 76 

2a. Crotalaria nitens HBK var. nitens Figure 2, Map 1. 

near Hydram, Rio Frio Augustine, Mt. Pine Ridge, Hunt 214 (us). 
GUATEMALA. VERAPAZ: Coban, 1,350 m., Türkheim 1948 (MICH, 
NY, US). HONDURAS. Comayagua, near Siguatepeque, 1,080-1,400 
m., Standley 56370 (Us). MEXICO. CHIAPAS: Muncipio of Tenejapa, 
6,000 ft, Breedlove 10976 (US); OAXACA: Entre Puerto Eligio a 
Comaltepec, Calderon 394 (MICH, US); VERACRUZ: Zacuapan, Purpus 
8007 (NY, US). 

2b. Crotalaria nitens HBK var. gracilis Windler, Phytologia 
21: 263, 1971. Figure 2, Map 1. 

Type: Mexico: Jalisco; Sierra de Halo, logging road 
7 miles south southwest of Tecalitlan and extending south- 
east toward San Isidaro, 13-16 miles from highway. Mc- 
Vaugh & Koelz 1188 (MICH). 

3. Crotalaria brevipedunculata Windler, Phytologia 21: 257. 

Shrub or suffrutescent herb with root type unknown. 
Stems more than 3 dm. long, 3.5 mm. thick, the longest 
internode 1.2 em. long; vestiture of dense, short spreading 
trichomes 0.6-0.7 mm. long. Stipules absent. Leaves elliptic 
or elliptic-oblong, 2.1-4.4 em. long, 5-13.5 mm. wide, base 
euneate to widely acute, the apex acuminate; vestiture of 
loosely appressed trichomes 0.5-0.9 mm. long; petioles 1.6- 
2.1 mm. long. Inflorescenses terminal and also leaf-opposed, 
peduncles 1.2-6.1 cm. long. Bracts sessile, linear or nar- 
rowly lanceolate, 4-4.4 mm. long, 0.3-0.6 mm, wide; pedicels 
3.8-4.2 mm. long. Calyces 10.5-11 mm. long, the tubes 2.5 
mm. long, bearing spreading trichomes 0.5-0.7 mm. long; 
bracteoles linear, 3.5 mm. long, 0.2-0.3 mm. wide. Corollas 
yellow, the standards 10.5-11 mm. long, equaling the upper 
calyx lobes; elongate anthers 1.6-2 mm. long, short anthers 
0.3-0.4 mm. long; style 6.4 mm. long. Fruits and seeds not 
known. Chromosome number: not known. Flowering date: 
December 20. Habitat: shady canyon slope with oaks and 
palms, elevation 3,500 feet. Range: Mexico: Durango, 
Sinaloa. Figure 3, Map 2. 
= Type: Sierra Tres Picos, Durango, infrequent, scattered. 
Gentry 5311 (GH). 

1974] Crotalarias — Windler 165 

Figure 3. Crotalaria brevipedunculata. 
A. Habit 
B. Stem 

166 Rhodora [Vol. 76 

X C.brevipedunculata | 

€ C.mexicana M LL—— — 
6 C. polyphylla 

e C.quercetorum 
——. | | 
Map 2. Distribution of Crotalaria brevipedunculata, C. mexicana, 
C. polyphylla, and C. quercetorum. 

Crotalaria brevipedunculata is characterized by its lack 
of stipules, its short terminal inflorescences and small 
flowers. It most nearly resembles C. nitens, but differs 
from it in its smaller flower size and shorter peduncles. 

In addition to the holotype only one other collection of 
this species has been observed: Mexico. SINALOA: Puerto 
a Tamiapa, Gentry 5815 (MICH, NY). 

4. Crotalaria polyphylla Riley, Kew Bulletin 43: 333. 1923. 

Perennial decumbent or ascending herb with a taproot 
to 1.4 em. thick. Stems 4-10, 1.2-3 dm. long, 1-2.5 mm. 
thick, the longest internode 1-3.2 cm, long; vestiture of 
short appressed or long spreading trichomes, the appressed 
0.3-0.6 mm. long, the spreading 0.8-1.2 mm. long. Stipules 

1974] Crotalarias — Windler 167 

Figure 4. Crotalaria polyphylla. 
A. Habit and leaf variation 
B. Stem 

168 Rhodora [Vol. 76 

absent. Leaves elliptic, oval, or elliptic-oblong, 1.6-4.2 em. 
long, 5-183 mm. wide, base rounded to cuneate, the apex 
obtuse, rounded, or acuminate; vestiture of both surfaces 
of loosely appressed trichomes 0.5-1.1 mm. long; petioles 
1-2.5 mm. long. Inflorescences terminal or terminal and also 
leaf-opposed, peduncles 4-16 em. long. Bracts sessile, nar- 
rowly elliptic to narrowly lanceolate, 2.5-9.5 mm. long, 
0.3-1.4 mm. wide; pedicels 4-6 mm. long, Calyces 9-13.5 
mm. long, the tubes 2-3 mm. long, bearing appressed tri- 
chomes 0.3-1 mm. long; bracteoles linear, narrowly lanceo- 
late or narrowly elliptic, 3-7 mm. long, 0.4-1 mm. wide. 
Corollas yellow, the standards 10-14 mm. long, 0.5-1.5 mm. 
longer than the upper calyx lobes; elongate anthers 1.7-2.3 
mm. long, short anthers 0.5-0.7 mm. long; styles 6-7.3 mm. 
long. Fruits 1.4-2.4 em. long, 0.5-0.9 em. broad; seeds green, 
grey-green, green-brown, brown, or tan, 7-18 per pod, 
1.8-2.5 mm. long. Chromosome number n—16. Flowering 
time: June-October. Habitat: slopes in open pine woods, 
elevation 6,500-9,500 feet, Range: Mexico: Chihuahua, 
Durango, Jalisco, Sinaloa. Figure 4, Map 2. 

Type: Mexico: Sierra Madre, Seeman 2187 (K!). 

Material here treated as Crotalaria polyphylla was in- 
cluded by Senn (1939) in C, maritima var. maritima; 
however, its terminal inflorescences and complete lack of 
stipules clearly separate it from C. rotundifolia (for treat- 
ment of Senn's C. maritima, see C. rotundifolia var. rotundi- 
folia). The epithet polyphylla was used by Senn as the 
varietal name for Mexican plants he considered to be C. 
purshii (C. purshii var. polyphylla). Personal examination 
of the type has led to the recognition of C. polyphylla at 
the specific level. (For treatment of plants previously 
placed in C. purshii var. polyphylla by Senn see C. querce- 
torum.) Crotalaria polyphylla is characterized by its 
decumbent or ascending habit, lack of stipules, and terminal 

Rio Mayo, pine slopes, Standley 2701 (GH, MEXU, US). DURANGO: 

1974] Crotalarias — Windler 169 

32.7 mi. W of Durango in allano slope below pine forest, 8,500 ft., 
Johnston 2694 (MEXU, MICH). JALISCO: Sierra de la Campana, 
along road to Mascota, 1,900-2,000 m., McVaugh 13777 (MEXU, MICH, 
US). SINALOA: Ocurahui, Sierra Surotato, Gentry 6227 (MICH, NY). 

5. Crotalaria stipularia Desvaux, J. Bot. (Desvaux) 3: 76. 

Crotalaria espadilla HBK. Nov. Gen. et Sp. Plantarum 
6: 399. 1824. Type: Humboldt 1025, (P; observed IDC 

Crotalaria stipularia var. serpyllifolia DC., Prodromus 
2: 124. 1825. Type: (G; observed DC Microfiche in Pro- 
dromus Herbarium). 

Crotalaria stipularis, orthographic error by Grisebach, 
Flora Brit. West Ind. Isl. 178. 1864. 

Crotalaria stipularis var. oblongata Grisebach, Flora 
Brit. West Ind. Isl. 178. 1864. 

Crotalaria stipularis var. sericea Grisebach, Flora Brit. 
West Ind. Isl. 178. 1864. 

Crotalaria stipularis var. espadilla Kuntze, Revisio 
Generum Plantarum 1: 175, 1891. 

Crotalaria stipularia forma eliptica Chodat & Hassler, 
4: 835. 1904. 

Crotalaria stipularia var. grandifolia Senn, Rhodora 41: 
333. 1939. Type: Hahn 239 (GH). 

Annual, erect herb with a taproot to 0.6 cm. thick. Stem 
solitary, 1-6.3 dm. long, 1.2-5 mm. thick, the longest inter- 
node 3-8 em. long; vestiture of loose appressed and/or 
spreading trichomes, 0.5-3 mm. long. Stipules present, 
decurrent the length of the internode, 1.1-3.2 cm. wide at 
the top, tapering to the subtending node, the stipule lobes 
ineurved 1.2-1.6 em. long. Leaves oval, oblong, elliptie, or 
lanceolate, 2.2-8.2 cm. long, 7.5-25 (-37) mm. wide, base 
rounded to cuneate, the apex retuse, rounded, acuminate, 
or acute; vestiture of both surfaces of appressed or loosely 
appressed trichomes 0.5-2.1 mm. long; petioles 0.5-1.5 mm. 
long. Inflorescences leaf-opposed, peduncles 3.1-10 cm. long. 
Bracts petiolate, elliptic, lanceolate to linear, the petioles 

170 Rhodora [Vol. 76 

Figure 5. Crotalaria stipularia. 
A. Habit and leaf variation 
B. Stem 

1974] Crotalarias — Windler 171 

1-1.6 mm. long, the blades 3-5 mm, long, 0.3-1.2 mm. wide; 
pedicels 2-4 mm. long. Calyces 8-10 mm, long, the tubes 
2.2.3 mm. long, bearing appressed or loosely appressed 
trichomes 0.7-1.2 mm. long; bracteoles elliptic, lanceolate, 
or linear, 3-4.5 mm. long, 0.4-0.7 mm. wide, Corollas yel- 
low, the standards 6-10 mm. long, equaling to 1.5 mm. 
shorter than the upper calyx lobes; elongate anthers 1.1-1.3 
mm. long, short anthers 0.3-0.5 mm. long; styles 4.5-5.2 
mm. long. Fruits 2.5-3.3 cm. long, width not known; seeds 
tan, bronze, red-brown, or dark brown, 17-41 per pod, 2-3 
mm. long. Chromosome number: n—16. Flowering time: 
February-December. Habitat: sandy soils, savannahs, 
clearings, fields and roadsides, elevations to 2,300 feet. 
Range: West Indies; Dominican Republic, Dominica, 
Guadeloupe, Haiti, Martinique, Puerto Rico, Trinidad. 
(Also widely distributed in South America.) Figure 5, 
Map 1. 
Type: Not located. 

Crotalaria stipularia, as treated by Senn, was divided 
into three varieties. Varieties "typica" and “orandifolia” 
are not different except in overall plant size. Both varia- 
tions are found in a single location mass collection from 
Puerto Rico sent to me by Dr. R. A. Howard of the Arnold 
Arboretum. This size variation is probably typical of plants 
of this species at most locations. Senn’s application of De 
Candolle’s C. stipularia var. serpyllifolia is to a plant in 
which all of the original leaves were lost during drought 
(or some other catastrophe) and new branches with small 
leaves and shorter internodes had developed before the 
plant was collected. The wide decurrent stipules were not 
lost when the leaves were lost, which accounts for the 
variety having large stipules. The IDC Microfiche photo- 
graph of De Candolle’s specimen in the Prodromus Her- 
barium shows a plant with smaller leaves, but not quite 
so small as those in Senn’s cited specimens. No field col- 
lections were made of C. stipularia during this study, but 
seeds were obtained for garden studies (Irwin 1050 and 
Stimson 4264). 

172 Rhodora [Vol. 76 

Crotalaria stipularia is characterized by its erect habit, 
leaf-opposed inflorescences, small flowers, and large stipules 
with incurved lobes. It most nearly resembles C. sagittalis, 
but the two taxa may be separated by stipule characters. 

855 (NY). DOMINICAN REPUBLIC. Prov. de la Vega: near Gara- 
bawa at Rio Yaquir, Fuertes 1669 (A, NY, P). GUADELOUPE. 
Montebello, alt. 50 m., Questel 432 (P, us). HAITI. Dept. du Nord: 
east of Marmelade, grassy summit of mountain, Leonard 7222 (us). 
MARTINIQUE. Fort Vaillant, Hahn 239 (GH; Photo., Us). PUERTO 
RICO. Along railroad N of Mayaguez, Heller 4574 (MICH, NY, US). 
ST. VINCENT. near Barrovollic (Leeward), Smith & Smith 1053 
(NY). TRINIDAD. south of Dabadie, Bulton & Hazen 702 (GH). 

6. Crotalaria sagittalis L., Sp. Pl. p, 714. 1753. 

Crotalaria fruticosa Miller, Gard. Dict. (pages not num- 
bered) 1768. Type: Jamaica, Houstoun, (BM, photo., NCU). 

Anonymos sagittalis (L.) Walt., Flora Carol. 181. 1788. 
Type: same as for C. sagittalis L. 

Crotalaria parviflora Roth, Catalecta Bot. 1: 83. 1797. 
Type: not located. 

Crotalaria sagittalis var. B oblonga Michx. Flora Boreali- 
Am. 2: 55. 1803. Type locality: "Virginia et Carolina". 

Crotalaria platycarpa Link, Enum. Pl. Hort. Reg. Bot. 
Berol. Alt. part 2, 227. 1822. Type: destroyed in Berlin. 

Crotalaria pringlei A. Gray, Proc. Am. Acad. 17: 200. 
1881-2. Type: Arizona, Pringle, summer 1881 (GH). 

Crotalaria sagittalis var. fruticosa (Miller) Fawcett and 
Rendle, Fl. Jamaica 4: 10. 1920. 

Crotalaria sagittalis var. blumeriana Senn, Rhodora 41: 
339. 1939. Type: Blumer 1772 (GH). 

Crotalaria tuerckheimii Senn, Rhodora 41: 339. 1939. 
Type: von Tuerckheim II 1282 (GH). 

Crotalaria belizensis Lundell, Amer. Midl, Naturalist 29: 
473. 1943. Type: Gentle 4144 at (NY). 

Annual or rarely perennial, erect herb with a taproot to 
0.6 cm. thick. Stems 1 (-7), 0.25-4.1 dm. long, 1-2.5 mm. 
thick, the longest internode 0.4-4.2 em. long; vestiture of 
long spreading trichomes, 1.5-3 mm. long. Stipules present, 

1974] Crotalarias — Windler his 

Figure 6. Crotalaria sagittalis. 
A. Habit and leaf variation 
B. Stem 

Rhodora [ Vol. 

Map 3. 

Tet "ese 
e! NA 


C. sagittalis 

- YT) = on 
- > J 
252525» u o 1 ka 
o i 
^ — 
} 297... 
W " —7» 

Distribution of Crotalaria sagittalis. 

1974] Crotalarias — Windler 175 

decurrent from 1% to the full length of the internode, 0.35- 
1.7 cm. wide at the top, tapering to the subtending node, the 
stipule lobes spreading 0.4-1.8 cm. long. Leaves elliptic, 
oblong, lanceolate, or linear-lanceolate, 1.1-10 cm. long, 
2.5-15 (22.5) mm. wide, base rounded to cuneate, the apex 
obtuse, mucronate, acuminate, or acute; vestiture of both 
surfaces of spreading or loosely appressed trichomes 1-2.9 
mm. long; petioles 1.2-2.5 mm. long. Inflorescences leaf- 
opposed, peduncles 2-8 (-12) cm. long. Bracts petiolate, 
lanceolate to elliptie lanceolate, the petioles 0.5-1.5 mm. 
long, the blades 4-8.1 mm. long, 0.5-1.3 mm. wide; pedicels 
3.5-5.5 mm. long. Calyces 7-15 mm. long, the tubes 2.3-3.9 
mm. long, bearing loosely appressed trichomes 0.8-2 mm. 
long; bracteoles elliptic lanceolate, lanceolate, or linear, 
2.5-7 mm. long, 0.7-1.2 mm. wide. Corollas yellow, the 
standards 6-12.5 mm. long, equaling to 4 mm. shorter 
than the upper calyx lobes; elongate anthers 0.9-2 mm. 
long, short anthers 0.3-0.5 mm. long; styles 4-5.3 mm, long. 
Fruits 1-2.7 (3.6) cm. long, 0.5-1.1 cm. broad; seeds tan, 
tan-green, olive, green-brown, or coffee-brown, 7-32 per pod, 
1.4-3 mm. long. Chromosome number: n=16. Flowering 
time: May-October. Habitat: road and railroad rights-of- 
way, disturbed ground, sand deposits, fields, prairies, lake 
margins, and dry oak and pine forests, elevation sea level 
— 8,500 feet. Range: Eastern and Central United States 
through Mexico and Central America to Panama and on 
the Greater Antilles. Figure 6, Map 3. 

Lectotype: “2 Kalm sagittalis" (LINN) (see Taxon 21: 

Type locality: “Brasilia, Virginia" as cited in Species 

Plants included here under Crotalaria sagittalis include 
material cited by Senn as variety blumeriana and variety 
fruticosa, as well as Senn’s variety "typica". Early leaves 
of plants of C. sagittalis tend to be broader than the later 
leaves, and all branches tend to have narrower leaves than 
the main stem. In addition, the further south a plant occurs, 
the longer the plant grows, and the more early leaves are 

176 Rhodora [Vol. 76 

lost before the end of the growing season. There is also 
a tendency for southern plants to have narrower early 
leaves than northern plants. Garden studies have shown 
Crotalaria sagittalis var. blumeriana to be a dry habitat 
ecophene of the typical species. Although a great deal of 
variation exists in C. sagittalis, especially in the Mexican 
representatives, little in the way of a clear pattern that 
would allow a useful segregation of the plants (and popu- 
lations) into varieties has been detected. Further study of 
this wide-spread but poorly understood species, to gain a 
more complete picture of the relationships between plant 
variation, geographic, and ecological factors, may prove 
helpful, especially in Mexico, Central and South America. 

Lundell 665 (vus). COSTA RICA. Paturage a San Jose de Costa 
Rica, Tonduz 443 (MicH, P). CUBA. Pinar del Rio: Herradura, 
Britton, Britton, Earle, & Gager 6404 (NY). GUATEMALA. Alta 
Verpaz: Coban, alt. 4,300 ft., Tuerckheim 239 (GH, Us). HAITI. 
Massif de la Selle Nouvelle Touraine, alt. 1,400 m., Ekman 1501 
(Ny, US). HONDURAS, Morazan, hills around Zamorano Valley, 
alt. 800-900 m., Swallen 11252, (us). JAMAICA. Castleton, 490 ft., 
Harris 11855 (GH, NY, P, US). NICARAGUA. Comarca ElCabo: 
LaTronquera, alt. 60 m., Molina 14881 (Ny). PANAMA. Panama: 
Sabana de Dormisolo, near Chepo, Pittier 4686 (Us). EL SALVA- 
DOR. Ahuachapan: Ahuachapan, alt. 800-1,000 m., Standley 19747 
(GH, NY, US). MEXICO. Baja California: S side of La Carrerita, 
Sierra El Taste, 1,500-1,600 m., Carter & Chisaka 8504 (MEXU). 
Chiapas: Tenejapa, 7,800 ft., Breedlove 19718 (us). Chihuahua: 
Madera, Pennell 19228 (vs). Colima: Colima, Palmer 130 (vs). 
Durango: without locality, Garcia 942 (vs). Guanajuato: NE of 
Guanajuato, Ripley & Barneby 13351 (NY). Jalisco: near Chapala, 
Rose & Painter 7636 (NY). Mexico: Puerto del Salitre, 1,300 m., 
Hinton 1780 (GH, MEXU, NY). Michoacan: vicinity of Coalcoman, 
1,000 m., Hinton 12862 (Ny, US). Morelos: Cuernavaca, Rose & 
Painter 6850 (GH, Ny, US), Nayarit: Cerro de la Cruz, E of Tepic, 
Mexia 669 (GH). Oaxaca: near city of Oaxaca, Rose & Hough 4623 
(NY, US). Puebla: 72 km. SE of Mexico City, Weaver 995 (US). 
San Luis Potosi: 7-8 mi. NE of Xilitla, King 4360 (us). Sinaloa: 
near Mazatlan, Windler & Windler 2885 (NcU). Sonora: NE cf El 
Tigre, 6,000 ft, White 4370 (GH, MICH). Tamaulipas: vicinity of 
Gomez Farias, ca. 350 m., Palmer 341 (us). Veracruz: SE of Cate- 
maco, meadow, Dressler & Jones 114 (MEXU, MICH, NY, US), UNITED 

1974] Crotalarias — Windler 177 

STATES. Alabama: MOBILE C0.: Dauphin Island, Jones 585 (ALA). 
Arkansas: SALINE CO.: ridges, Benton, Demaree 37333 (stu). Con- 
necticut: MIDDLESEX CO.: Middlefield, Kofoid (GH). Georgia: JACK- 
SON CO.: 11 mi. N of Athens, Cronquist 4571 (GH). Illinois: JACKSON 
co.: roadside S of Elkville, Bailey & Swayne 462 (stu, NCU). Indiana: 
LAKE CO.: East Gary, Hull 18 July 1936 (GH). Iowa: JONES CO.: 
railroad right-of way, Sect. 7, Oxford Twp., Isely 5913 (GA). Kansas: 
RILEY Co.: ravines, Norton 80 (GH, P). Kentucky: GRAVES CO.: rail- 
road embankment, Fancy Farm, O'Dell & Windler 936 (stv). Louis- 
iana: CALCASIEU PARISH: 114 miles NNW of Starks, Sect. 26, Las- 
seigne 1579 (LAF). Maryland: Montgomery?, Wildwood, Dowell 1626 
(GH). Massachusetts: MIDDLESEX CO.: shore of Winter Pond, Bartlett 
224 (GA, GH, TEX, WIS). Michigan: KALAMAZOO CO.: 1 mi. E of 
Schoolcraft, Hanes 1796 (GH). Minnesota: CHISAGO CO.: Lindstrom, 
Anderson 6458 (GH, WIS). Mississippi: LAYAYETTE CO.: near Tula 
Cemetery, 18 mi. SE of Oxford, Pullen 64294 (GA, MISS, NCU). 
Missouri: BARTON CO.: prairies, Milford, elev. 750 ft., Demaree 39377 
(GA, NCU, USF). New Jersey: BERGEN CO.: Hackensack, route 4, 
sandy field, Rush & Svenson 6769 (GH, wis). New York: SARATOGA 
co.: N of Crescent, Burnham 8 Sept. 1906 (GH). North Carolina: 
CALDWELL CO.: 1 mi. SE of Hudson, Randolph & Randolph 1096 
(GH, NCU). Oklahoma: MUSKOGEE CO.: 2 mi. S of Braggs, Edmond 
164 (TEx). Pennsylvania: DELAWARE CO.: Wayne, dry gravel bank, 
Bartram 1162 (GH). Rhode Island: Cumberland (?) without collec- 
tion name & number, Aug. 1840 (GH). South Carolina: GREENVILLE 
CO.: 215 mi. SW of Berry's Pond, Freeman 56577A (NCU). Tennessee: 
KNOX CO.: sandy soil near Knoxville, Ruth 661 (wis). Texas: JASPER 
co.: 3 mi. N of Evadale, Turner & Tharp 3079 (TEX). Vermont: 
WINDHAM CO.: Vernon, Blanchard 1901 (GH). Virginia: CULPEPER 
co.: 0.3 mi. N of Lakota on Co. Rt. 621, Ahles & James 61692 (NCU). 
West Virginia: HARDY co.: Lost River Bridge, wasteground, Core 
18 Aug. 1931 (GH). 

7. Crotalaria purshii DC., Prodromus 2: 124. 1825. 
Crotalaria sagittalis var. B L. Sp. Pl. 714. 1753. 
Crotalaria sagittalis var. a linearis Michx., Fl. Boreali- 

Am. 2: 55. 1808. Type locality: “Virginia et Carolina". 
Crotalaria laevigata Pursh, Fl. Am. Sept. 469, non Lam., 

Encyc., 2: 198. 1786. Type: Pursh specimen not located ; 

Syntype was plate in Plukenet, Phytographia-Amagestum 

Bot. t. 277, fig. 2. 1694. Specimen from which this figure 

was drawn is in Sloane collection at BM. 

Perennial, erect herb with a taproot to 0.7 cm. thick. 

178 Rhodora [Vol. 76 

Figure 7. Crotalaria purshii. 
A. Habit and leaf variation 
B. Stem 

Crotalarias — Windler 

C. purshii X 

Map 4. Distribution of Crotalaria purshii. 

HAA A as ae 
Pe HO UI i Aro - C 
: VETE CHI Lad Re 


180 Rhodora [Vol. 76 

Stems 1-4 (8), 1.5-5 dm. long, 1-1.5 mm. thick, the longest 
internode 3.3-8.5 cm. long; vestiture of short appressed or 
loosely appressed trichomes 0.4-0.8 mm. long. Stipules 
present, decurrent one-half to the full length of the inter- 
node, 0.3-0.9 cm. wide at the top tapering to the base, the 
stipule lobes spreading 0.5-1.7 cm. long. Leaves linear, 
oblong, linear-lanceolate or elliptic, 4.9-10 cm. long, 1.7-12 
mm. wide, base rounded to cuneate, the apex obtuse or 
acute; the lower surface bearing appressed trichomes 0.3- 
0.7 mm. long, the upper surface glabrous; petioles 0.5-2 
mm. long. Inflorescences leaf-opposed, peduncles 7.5-23 cm. 
long. Bracts petiolate, elliptic-lanceolate to lanceolate, the 
petioles 0.5-1.6 mm. long, the blades 4.5-7 (8.5) mm. long, 
0.6-1.8 (2) mm. wide; pedicels 4.5-6 mm. long. Calyces 
9-12.65 mm. long, the tubes 2.5-3.5 mm. long, bearing ap- 
pressed trichomes 0.2-0.5 mm. long; bracteoles lanceolate 
or ellipite-lanceolate, 3.5-5 mm. long, 0.8-1.2 mm. wide. 
Corollas yellow, the standards 8-13.5 mm. long, 2 mm. 
shorter to 0.5 mm. longer than the upper calyx lobes; 
elongate anthers 1.2-1.6 mm. long, short anthers 0.3-0.6 
mm. long; styles 2.7-4.6 mm. long. Fruits 1.4-2.8 cm. long, 
0.6-1 cm. broad; seeds tan, olive, green-brown, or dark 
brown, 11-37 per pod, 1.8-2.9 mm. long. Chromosome num- 
ber: n=16. Flowering time: March-October. Habitat: 
savannahs, sandhills, and open pine woods, coastal plain. 
Range: United States: Alabama, Florida, Georgia, Louis- 
iana, Mississippi, North Carolina, South Carolina, Virginia. 
Figure 7, Map 4. 
Type: Based on Crotalaria laevigata Pursh. 
Type locality: Pine woods of Virginia and Carolina. 

Crotalaria purshii, named by De Candolle, was based on 
material from Virginia and Carolina. He cites C. laevigata 
of Pursh as being the same, but proposes the name C. 
purshii since C. laevigata had been previously used by 
Lamarck. Crotalaria sagittalis var. B L. is based on 
Plukenet's figure (Plukenet, 1694, t. 277, Fig. 2) which 
is purshii. 

Senn (1939) recognized a variety, polyphylla, under 

1974] Crotalarias — Windler 181 

Crotalaria purshii. Most specimens of this variety are here 
treated as C. quercetorum. For further information con- 
cerning the epithet polyphylla see the discussion under C. 

Crotalaria purshii is characterized by its erect habit; 
sparse, short, appressed pubescence; spreading stipule 
lobes; frequently narrow leaves, the upper surfaces of which 
are glabrous; and long peduncles. Crotalaria purshii is 
similar to C. sagittalis in habit and general appearance in 
parts of its range, but it may be distinguished from C. 
sagittalis by its glabrous upper leaf surfaces, appressed 
pubescence, and long peduncles. The extremely variable 
C. rotundifolia var. rotundifolia may occasionally bear a 
resemblance to C. purshii, but these plants lack glabrous 
upper leaf surfaces. 

WIN co.: sandy field, Jones 496 (aua). Florida: CALHOUN CO.: 
Cypress Park, Dead Lakes, Godfrey & Clewell 62916 (LAF). Georgia: 
WAYNE CO.: open pinelands, 5 mi. S. of Jessup, Duncan 6941 (GA). 
Louisiana: ST. TAMMANY PARISH: 7% mi. S of town of Pearl River, 
Sect. 13, Lasseigne 1480 (LAF). Mississippi: PEARL RIVER CO.: 1 mile 
N of Picayune along Hwy. 12, low pine lands, Bollwinkel & Wun- 
derlin 9 (stu). North Carolina: CRAVEN CO.: pine savannah, 0.6 mi. 
SW of Havelock, Radford 34504 (NcU). Virginia: ISLE OF WIGHT CO.: 
dry sandy pine and oak woods ca. 1 mile SE of Zuni, Fernald & 
Long 6233 (GH). 

8. Crotalaria quercetorum Brandegee, U. Cal. Pub. Bot. 
10: 407. 1924. 

Crotalaria purshii var. polyphylla (Riley) Senn, Rhodora 
41: 346. 1939. (see note below) 

Annual, erect, or ascending herb with a slender taproot 
to 0.25 em. thick. Stem solitary, 1.1-5.3 dm. long, 1-3.1 mm. 
thick, the longest internode 1.3-5.2 cm. long; vestiture of 
short appressed trichomes, 0.3-0.4 mm. long. Stiples usually 
absent or very narrow, decurrent 1/5 the length of the 
internode, 0.3-0.4 cm. wide at the top narrowing to the 
base, the stipule lobes spreading 0.1-0.25 cm. long. Leaves 
elliptic, narrowly oblong, narrowly elliptic, narrowly lanceo- 

182 Rhodora [Vol. 76 

Figure 8. Crotalaria quercetorum. 
A. Habit and leaf variation 
B. Stem 

1974] Crotalarias — Windler 183 

late, or linear, 3-6.8 cm. long, 2-5.5 (-8) mm. wide, base 
rounded to cuneate, the apex acute or acuminate; vestiture 
of appressed trichomes 0.2-0.5 mm. long, the upper surface 
usually glabrous; petioles 1.5-3 mm. long. Inflorescences 
leaf-opposed, peduncles 2.5-8 cm. long. Bracts sessile, nar- 
rowly elliptic, or linear, 2.5-3.5 mm. long, 0.2-0.3 mm. wide; 
pedicels 2.5-4.2 mm. long. Calyces 8.2-11 mm. long, the 
tubes 1.5-2 mm. long, bearing appressed trichomes 0.2-0.8 
mm. long; bracteoles linear or narrowly elliptic, 1.5-3.5 mm. 
long, 0.2-0.3 mm. wide. Corollas yellow, the standards 
7.5-10.5 mm. long, equaling to 2 mm. shorter than the 
upper calyx lobes; elongate anthers 0.8-1.4 mm. long, 
short anthers 0.2-0.3 mm. long; styles 3.5-3.6 mm. long. 
Fruits 1-2.4 cm. long, 0.4-0.7 cm. broad; seeds olive, green- 
brown, or brown, 24-30 per pod, 1.4-2 mm. long. Chromo- 
some number: n—16. Flowering time: August-March. 
Habitat: ridges and slopes in oak, oak-pine, and pine woods, 
elevation 2,600-7,000 feet. Range: Mexico: Chiapas, 
Jalisco, Michoacan, Nayarit, Oaxaca, Sinaloa. Figure 8. 
Map 2. 

Type: Mexico: Chiapas: Hacienda Monserrate, Purpus 
9144 (uc!). 

Crotalaria quercetorum was included under C. purshii 
by Senn. Unfortunately, Senn chose to use the epithet 
polyphylla for what he recognized as a variety of C. purshit. 
Examination of the type of C. polyphylla Riley leads to the 
conclusion that the name is erroneously applied to the ma- 
terial here treated as C. quercetorum. The holotype of 
C. quercetorum is a mixed sheet with one specimen present 
which may be interpreted as intermediate between C. 
quercetorum and C. sagittalis. The isotype (US) examined 
is also composed of several elements, but all are referable 
to C. quercetorum. 

Crotalaria quercetorum is characterized by its short 
appressed pubescence, lack or near lack or stipules, usually 
narrow foliage, short peduncles and small flowers. Crota- 
laria quercetorum is similar to C. purshii but may be dis- 
tinguished by its much shorter peduncles and its usually 

184 Rhodora [Vol. 76 

smaller size. Because a number of specimens with inter- 
mediate morphology have been collected, hybridization 
between C. quercetorum and C. sagittalis is suspected in 
parts of their range where they are sympatric. Windler 
& Windler 2994 (NCU) is a collection in which some speci- 
mens seem to reflect this putative hybridization. 

given, Bernoulli 282 (G). MEXICO. cnurAPAS: Hacienda Monser- 
rate, Purpus 9144 (uc, Us); Jalisco: 15-20 mi. SE of Autlan-Sierra 
Manantlan ca. 1,700 ft., McVaugh 13973 (MICH); Michoacan: 32 mi. 
W of Jiquilpan along road to Manzanillo, 7,000 ft, Fearing 1195 
(TEX); Nayarit: 1 mi. from town of Santa Maria del Oro on way 
to Laguna Santa Maria, 3,700 ft., Windler & Windler 2917 (NCU); 
Oaxaca: 5-6 Km NE of Putla, 350 m., McVaugh 22248 (MICH); 
Sinaloa: Puerto a Tamiapa, 4,500 ft., Gentry 5815 (MEXU). 

9. Crotalaria mexicana Windler, Phytologia 21: 259. 1971. 
Crotalaria sagittalis var. fruticosa (Miller) Fawcett and 
Rendle, 4: 10, 1920, pro parte, sensu Senn, non sensu typus. 

Annual erect herb with a slender taproot to 0.3 cm. thick. 
Stems solitary, 1.2-2.3 dm. long, 1.6-2.4 mm. thick, the 
longest internode 1-1.5 em. long; vestiture of dense, ap- 
pressed trichomes, 1.2-2.5 mm. long. Stipules absent. 
Leaves narrowly elliptic, linear-lanceolate or linear, 2.2-4.6 
cm. long, 4-8 mm. wide, base rounded to cuneate, the apex 
rounded, acute or acuminate; vestiture of both surfaces of 
dense, loosly appressed trichomes 1.1-2.1 mm. long; petioles 
0.5-0.6 mm. long. Inflorescences leaf-opposed, peduncles 
0.8-2 cm. long. Bracts sessile, lanceolate, 3.3-3.6 mm. long, 
0.7-0.8 mm. wide; pedicels 2.5-4 mm. long. Calyces 10.5-11.5 
mm. long, the tubes 2-2.5 mm. long, bearing loosely ap- 
pressed and spreading trichomes 0.8-2 mm. long ; bracteoles 
linear-lanceolate, 4-4.5 mm. long, 0.5-0.6 mm. wide. Corollas 
yellow, the standards 10-10.5 mm. long, equaling to 0.5 mm. 
shorter than the upper calyx lobes; elongate anthers 1.5-1.7 
mm. long, short anthers 0.5-0.6 mm. long; styles 4.8-5.3 
mm. long. Fruits 2.1-2.5 em. long, 0.8-1 cm. wide; seeds 
brown, number per pod not known, 1.8-2 mm. long. 
Chromosome number: not known. Flowering time: Sep- 

1974] Crotalarias — Windler 185 


EREE eee, 

= ee er te 



Figure 9. Crotalaria mexicana. 
A. Habit and leaf variation 
B. Stem 
C. Leaf pubescence 

186 Rhodora [Vol. 76 

tember-October. Habitat: dry slopes of mountains, eleva- 
tion ca. 6,000 ft. Range: Mexico: Jalisco. Figure 9, 
Map 2. 

Type: Mexico: Jalisco, mountainside above Etzatlan, 
Pringle 8855 (holotype, GH; isotypes, TEX, US). 

Crotalaria mexicana is a new species, the representatives 
of which were referred by Senn (1939) to C. sagittalis 
var. fruiticosa (here treated as C. sagittalis var. sagittalis). 
Crotalaria mexicana is most similar to C. sagittalis and 
C. quercetorum. It differs from C. sagittalis in its lack of 
stipules and absence of spreading pubescence, and from 
C. quercetorum in its short thick peduncles and in its 
dense pubescence. Crotalaria mexicana is characterized by 
its erect habit, dense, appressed pubescence, lack of stipules, 
and extremely short, leaf-opposed peduncles. 

Pringle 8855 (—type), Pringle 11807 (GH, US), Rose & Painter 7571 
(US); near Guadalajara, Rose & Painter 7469 (US). 

10. Crotalaria rotundifolia (Walt.) Gmelin in Linne . 
Systema Naturae 2: 1095. 1792. 

Anonymos rotundifolia Walter, Flora Caroliniana, 181. 
1788. Type: BM, photo., NCU! 

Crotalaria sagittalis var. ovalis Michx. Flora Boreali-Am. 
2: 55. 1803. Type locality: “Virginia et Carolina". 

Crotalaria rotundifolia Poiret, Encycl. Meth. 2: 402. 
1812, non Gmelin, 1792. (see Ward, 1962). 

Crotalaria ovalis (Michx.) Pursh, 469. 1814, nomen 

Crotalaria hookeriana A. DC. in A.P. & A. DC., Mem. 
Soc. Phys. Genéve 9: 97. 1841. Type: not observed. 

Crotalaria leptoclona Schauer, Linnaea 20: 737. 1847. 
Type: Mexico, Aschenborn 172, not located. 

Crotalaria maritima Chapman, Flora South. United 
States, Suppl. 614. 1883. Type: not located. 

Crotalaria linaria Small, Flora SE United States 679. 
1933. Type: Pine Key, Fla., Blodgett (NY!). 

1974] Crotalarias — Windler 187 

Crotalaria maritima var. linaria (Small) Senn, Rhodora 
41: 347. 1939. 

Crotalaria rotundifolia var. linaria (Small) Fernald & 
Schubert, Rhodora 50: 208. 1948. 

Perennial procumbent, decumbent, or ascending herb 
with a taproot to 1.2 cm. thick. Stems 2-20, 0.3-6.5 dm. 
long, 0.5-3.1 mm. thick, the longest internode 0.6-4.8 cm. 
long; vestiture of appressed, loosely appressed, or spreading 
trichomes, appressed 0.3-0.6 mm. long, spreading 0.8-3.2 
mm. long. Stipules present, with only the stipule lobes 
present or the stipules decurrent from 1, to the full length 
of the internode, 0.13-2.3 cm. wide at the top tapering to 
the base, the stipule lobes spreading 0.15-1.7 cm. long. 
Leaves round, oval, oblong, ovate, lanceolate, elliptic, or 
linear, 0.9-5.5 em. long, 1.0-33.1 mm. wide, base rounded to 
widely cuneate, the apex rounded, obtuse, mucronate, acu- 
minate, or acute; vestiture of both surfaces of erect, ap- 
pressed, or loosely appressed trichomes 0.3-2.8 mm. long; 
petioles 0.5-3 mm. long. Inflorescences leaf-opposed, pe- 
duncles 1.3-33.9 cm. long. Bracts with or without petioles, 
elliptic, lanceolate, or linear, the petioles to 1.5 mm. long, 
the blade 2-7.1 mm. long, 0.3-2 mm. wide; pedicel 2-5 mm. 
long. Calyces 7-14.9 mm. long; the tubes 1.5-4.5 mm. long, 
bearing appressed, loosely appressed or spreading trichomes 
0.2-1.5 mm. long; bracteoles broadly to narrowly elliptic, 
lanceolate, or linear, 1.8-6.5 mm. long, 0.3-1.3 mm. wide. 
Corollas yellow, the standards 6-15 mm. long, 2.5 mm. 
shorter to 1.9 mm, longer than the upper calyx lobes; 
elongate anthers 0.9-2.4 mm. long, short anthers 0.3-0.7 
mm. long; styles 4-6.9 mm. long. Fruits 1.2-2.8 cm. long, 
0.5-1.1 cm. broad; seeds tan, bronze, green-tan, olive, green- 
brown, or red-brown, 12-46 per pod, 1.5-2.7 mm. long. 
Chromosome number: n—16. Flowering time: United 
States, January-October; Mexico, July-March. Habitat: 
road and railroad rights-of-way, rocky slopes, sandridges, 
sandhills, open pine, and pine-hardwood forests, elevation 
sea level — 8,250 feet. Range: Costa Rica; Guatemala; 
Mexico: Chihuahua, Guanajuato, Hidalgo, Jalisco, Mexico, 

188 Rhodora [Vol. 76 

Michoacan, Morelos, Nayarit, Oaxaca, Puebla, San Luis 
Potosi, Sonora, Veracruz; El Salvador; United States: 
Alabama, Florida, Georgia, Louisiana, Mississippi, North 
Carolina, South Carolina, Virginia. 

Type: In Walter Herbarium at BM, photo observed. 

Crotalaria rotundifolia was first recognized as a distinct 
species by Walter (1788) who also recognized it as being 
congeneric with C. sagittalis L. Walter apparently thought 
that C. rotundifolia and C. sagittalis represented a new 
genus distinct from Crotalaria, but, instead of describing 
a new genus, he placed them in a miscellaneous category 
under the genus name Anonymos. A few years later, Gme- 
lin placed Walter's Anonymos species under other genera 
(Sprague, 1939). Anonymos rotundifolia was placed with 
C. sagittalis L. under Crotalaria. In 1812, Poiret again 
published the combination, and he, rather than Gmelin, is 
frequently attributed the authorship of the combination 
C. rotundifolia. In 1814, Pursh also placed Walter’s 
Anonymos rotundifolia into Crotalaria. He, however, pro- 
posed a new species name, C. ovalis, and cited Walter’s 
name as a synonym, making C. ovalis a superfluous epithet 
when published and therefore illegitimate. 

Senn (1939), after examining a photograph of Miller's 
type of Crotalaria angulata, thought that it was synony- 
mous with C. rotundifolia, and the name has since been 
used for the American plants. In 1897, Britten and Baker 
examined the specimen and concluded it did not differ 
from C. biflora L. A comparison of the photograph of 
Miller’s specimen with an IDC microfiche of a C. biflora 
specimen in the Linnaean Herbarium leads me to agree 
with Britten and Baker. In addition, Miller indicated that 
the flowers were produced singly on the sides of the 
branches and that the leaves were sessile. Neither of these 
characters agrees with the type from the Miller Herbarium. 
It may also be possible that Miller made a mistake in indi- 
cating the location of the place of origin of the seed. It 
now seems better to recognize the epithet of Walter and to 
reject the Miller name for our American plants. 

1974] Crotalarias — Windler 189 

Fernald and Schubert (1948), after studying photo- 
graphs of Walter's Herbarium (taken by Schubert) at the 
British Museum of Natural History, suggested that the 
name Crotalaria rotundifolia actually refers to the plants 
with short appressed pubescence previously referred to as 
C. maritima. A more recent photograph of the specimen 
figured by Fernald and Schubert has been supplied by Dr. 
Norman K. B. Robson of the British Museum. Dr. Robson 
(personal communication) indicates that the plant has 
“appressed-sericeous” pubescence and would “run down 
easily in Senn's key to C. maritima Chap.". In light of this 
information, the name C. rotundifolia is recognized as ap- 
plying to the material previously recognized as C. maritima, 
as Fernald and Schubert suggested. 

Crotalaria rotundifolia may be distinguished by its pro- 
cumbent, decumbent, or occasionally ascending habit; leaf- 
opposed inflorescences; and the presence of stipules. It is 
an extremely variable species, especially in the Florida 

Two varieties may be distinguished in the following 
Plants with appressed stem pubescence. .............. 
RONDE n 10a. C. rotundifolia var. rotundifolia. 
Plants with spreading stem pubescence, .............. 
10b. C. rotundifolia var. vulgaris. 

10a. Crotalaria rotundifolia var. rotundifolia Figure 10, 
Map 5. 

While plants of some populations in the higher Florida 
Keys and along the Florida Gulf coast have only narrow 
leaves, plants at most locations in the state are extremely 
variable. Crotalaria linaria Small was based on narrow 
leaved plants and was maintained by Senn as a variety of 
C. maritima. Since a narrow leaved plant usually cannot 
be differentiated as being from either the entirely narrow 
leaved populations or the heterogeneous populations on the 
peninsula, no purpose is served by maintaining the variety 
or by reducing it to the status of forma. 

190 Rhodora 

Figure 10. Crotalaria rotundifolia var. rotundifolia. 
A., B. Habit and leaf variation 
C. Stem 
Crotalaria rotundifolia var. vulgaris 
D. Habit and leaf variation 
E. Stem 

1974] Crotalarias — Windler 191 

C. rotundifolia 

Map 5. Distribution of Crotalaria rotundifolia, var. rotundifolia. 

[Vol. 76 



C. rotundifolia var vulgaris 

Map 6. Distribution of Crotalaria rotundifolia var. vulgaris. 

1974] Crotalarias — Windler 193 

white sand, Britton, Britton, & Brown 6653 (US). UNITED STATES. 
Florida: LEVY CO.: evergreen scrub oak sandridge, Cedar Key, 
Godfrey 56571 (GA, GH, USF). MONROE CO.: open dry sandy swale 
behind low coastal shell ridge, Middle Cape, Cape Sable, Everglades 
National Park, Ward 3939 (GA, LAF, NCU). Georgia: MCINTOSH CO.: 
sandhill thicket 1.3 mi. N of Ft. Barrington, Cox Road on ridge, 
Boseman 458 (NCU). Mississippi: JACKSON C0.: Ocean Springs 
and vicinity of Gulf Coast Research Laboratory, sandy soil under 
pines, west end of Deer Island, Channell 414 (usF). South Carolina: 
GEORGETOWN CO.: roadsides, 5.6 mi. S of Maryville, Radford 28670 

10b. Crotalaria rotundifolia var. vulgaris Windler, Phyto- 
logia 21: 264. 1971. 

Crotalaria angulata Miller, 1768, sensu Senn, 1939. (See 
discussion above. 

Figure 10, Map 6. 

Type: South Carolina: Hampton County, about three 
miles northwest of Yemassee on South Carolina Highway 
68. Sandhill. 23 July 1967, D. R. & B, K. Windler 2769 

Crotalaria rotundifolia var. vulgaris is distinguished by 
its spreading pubescence. Over most of its range var. 
vulgaris is also characterized by round to oval leaves, but 
in northern Florida and southern Georgia it intergrades 
with the usually narrower leaved var. rotundifolia. 

ft., Cooper 165 (us). GUATEMALA. Chimaltenango: alt. 2,100 m., 
Standley 61485 (NY). PANAMA. Panama: roadside, Rio Tapia, 
Standley 28214 (us). EL SALVADOR. vicinity of San Salvador, 
alt. 650-850 m., Standley 22444 (NY). MEXICO. Baja California: 
San Antonio, Cape region, Purpus 475 (US). Guanajuato: 20 mi. 
NW of Irapuato, Barkley, Rowel, & Parson 752 (TEx). Hidalgo: 
Dist. Zimapan, on road from Hwy. at km. 242 to Encarnacion, Moore 
& Wood 4319 (MEX). Jalisco: 5 mi. W of Guadalajara on Hwy. 15, 
beneath cliff face in road cut, brown sandy soil, Irwin 1257 (TEX). 
Mexico: Lecheria, Pringle 13411 (GH, US). Michoacan: growing in 
oak, 2 mi. above Tancitaro, elev. 7,000 ft, Leavenworth 517 (GH). 
Morelos: km. 56-57, road from D.F. to Cuernavaca, roadside em- 
bankment, Moore 125 (GH). Nayarit: 13 mi. SE of Tepic, in red 
clay, Irwin 1293 (TEX). Oaxaca: Valley of Oaxaca, alt. 5,500-7,500 

194 Rhodora [Vol. 76 

ft., Nelson 1481 (GH). Puebla: vicinity of Puebla, Arsene 1414 (Us). 
San Luis Potosi: 15 mi. E of Ciudad del Marz on Rt. 80, ca. 1,000 
m., Mickel 544 (MICH). Sinaloa: dry hill, vicinity of Mazatlan, 
Rose, Standley, & Russell 13823 (US). Sonora: 3 mi. NE of Matape, 
Wiggins & Rollins 400 (GH). Veracruz: Misantla, Purpus 5907 
(GH, NY, US). UNITED STATES. Alabama: CLARKE CO.: near 
Choctaw Lake, elev. 170 ft., Demaree 50694A (LAF, NCU). Florida: 
HERNANDO CO.: Chinsegut Hill Bird Sanctury, open area along 
margins of pine woods, Windler 1136 (NCU). Georgia: BEN HILL CO.: 
open pine-oak on sandy upland, 7.1 mi. N of Fitzgerald, Duncan & 
Hardin 14301 (GA). Louisiana: WASHINGTON PARISH: mixed woods, 
edge, along Pushepetappa Creek, 2 mi. WNW of Varnado, Thieret 
25910 (LAF). Mississippi: LAMAR C0.: 12 mi. WSW of Hattiesburg 
along Black Creek, low area, Doughty 162 (MISS). North Carolina: 
BLADEN CO.: wet ditch on 701 to White Lake, Radford & Radford 
2065 (NcU). South Carolina: BERKELEY CO.: common on sandy bank 
ea. 15 mi. S of Moncks Corner on 52, Wilbur 3547 (GA). Virginia: 
NANSEMOND CO.: dry white sand of pineland SW of Marsh Hill 
School S of South Quay, Fernald & Long 11050 (GH). 

11. Crotalaria bupleurifolia Schlecht. & Cham., Linnaea 
5: 575. 1830. 

Crotalaria heldiana A. DC. in A. & A.P. DC. Mem. Soc. 
Phys. Geneve 9: 97. 1841. Type: Grown from seed of un- 
known source in the garden at Carlsruhe, (G!). 

Annual or perennial spreading suffrutescent herb with 
a taproot to 2 cm. thick. Stems few-many, 5-27 dm. long, 
2.3.5 mm. thick, the longest internode 3.5-10 cm. long; 
vestiture of spreading or loosely appressed trichomes 0.2- 
1.3 mm. long. Stipules present, decurrent for 14, to the 
full length of the internode, 0.5-2.7 cm. wide at the top 
tapering slightly to or past the subtending node, the stipule 
lobes absent or spreading 0.5-3 cm. long. Leaves oval, ellip- 
tie, ovate, broadly lanceolate, or lanceolate, 3.2-10.2 cm. 
long, 7-46 mm. wide, base obtuse to cuneate, the apex 
obtuse, mucronate, acuminate, or acute; vestiture of both 
sides of spreading or loosely appressed trichomes 0.2-1.6 
mm. long or nearly glabrous; petioles 2-3 mm. long. Inflor- 
escences leaf-opposed, peduncles 2-16 cm. long. Bracts 
sessile, linear, subulate, or narrowly lanceolate, 4.5-7.5 mm. 
long, 0.3-0.6 mm. wide; pedicels 4-5.5 mm. long. Calyces 

1974] Crotalarias — Windler 195 
13.5-19 mm. long, the tubes 2.5-5 mm. long, bearing 
loosely appressed or spreading trichomes 0.1-1.1 mm. long; 
bracteoles linear, subulate, or narrowly lanceolate, 3-6 mm. 
long, 0.2-0.7 mm. wide. Corollas yellow, the standards 
16-25 mm. long, equaling to 6 mm. longer than the upper 
calyx lobes; elongate anthers 2-2.8 mm. long, short anthers 
0.5-0.7 mm. long; styles 7-11 mm, long. Fruits 3-4.1 cm. 
long, breadth not known; seeds tan, green-brown, or brown, 
number per pod not known, 2.2-2.5 mm. long. Chromosome 
number: »—16. Flowering time: August-March. Habitat: 
North-facing slopes and bluffs and cloud forests with firs 
and broad leaf trees, elevation 2,500-7,500 feet. Range: 
Mexico: Chiapas, Guerrero, Jalisco, Mexico, Michoacan, 
Sinaloa, Veracruz. 

Type: Hacienda de la Laguna? Schiede et Deppe s.n. 

Crotalaria bupleurifolia was described by Schlechtendal 
and Chamisso in an article describing the collections of 
Schiede and Deppe in Mexico. A specimen collected by 
Schiede and Deppe at G is marked “Typus” and may well 
represent the type even though the location on the label 
reals *Mexique. Cuerta grande de Chiconquiaco" rather 
than “Hacienda de la Laguna" cited with the original 
description of C. bupleurifolia. The number 596 cited with 
Schiede's name in Senn (1939) refers to the species number 
which accompanied Schlechtendal et Chamisso’s description. 

Crotalaria bupleurifolia is characterized by its generally 
large size, unusual stipules, and large flowers and habit. 
It most closely resembles C. nayaritensis but may be dis- 
tinguished from it by the large flower size, larger overall 
size, and usually wider stipules. 

Two varieties may be recognized: 

Stipules present only at the base of peduncles, decurrent 
for only a single mnternodes ys a es sete ee 
ee es LAM 11a. C. bupleurifolia var. bupleurifolia. 

Stipules present at the base of most leaves, frequently 
decurrent for more than one internode. ............ 

11b. C. bupleurifolia var. robusta. 

196 Rhodora [Vol. 76 

Figure 11. Crotalaria bupleurifolia var. bupleurifolia. 
A. Habit and leaf variation 
B. Stem 

Crotalarias — Windler 


\ ? 
` 2 
| ^ 
| x K 
éinioleh b diina 
E: ho i x 
H ) 
Lor mi 
. r 
z a3 
g i ] SA e © 
z PEERS m t VA e > 
LOJ g 
hee v Ec: . 
'e o 
S t. = 
[d E — 
3 3s 
$ i / 9 9 
a a 
: @ 3 3 
uU A A 2m 
9. — E / rr c3 
5 e GO 
te fi 
JS TU m——— 
2 | e 

© C.nayaritensis 



Map. 7. Distribution of Crotalaria bupleurifolia var. bupleurifolia, 

C. bupleurifolia var. robusta, and C. nayaritensis. 

198 Rhodora [Vol. 76 

lla. Crotalaria bupleurifolia var. bupleurifolia Figure 11. 
Map 7. 

Miranda 5746 (MEXU). Guerrero: San Antonio Buenos Aires, 
Hinton 11708 (GH, TEX, US). Jalisco: near Santa Monica, 1,950-2,050 
m, McVaugh 14086 (MEXU, MICH, US). Mexico: Rincon del Carmen, 
Dist. Tamascaltepec, 1,340 m, Hinton 1730 (MEXU, NY, US). Michoacan: 
NW of Aguililla, 15 Km S of Aserradero Dos Aguas, 1,650-1,700 m, 
McVaugh 22767 (MICH). San Luis Potosi: Las Canoas, Pringle 8 
(MEXU). Sinaloa: Batel, Concordia, Dehesa 1619 (US). Tamaulipas: 
10 Km NW of El Progresso, 1,450 m, Stanford, Retherford, & North- 
craft 1009 (GH). Veracruz: Barranca de Tenampa, Zacuapan, Pur- 
pus 3663 (GH, NY, US). 

llb. Crotalaria bupleurifolia var. robusta (Senn) Windler, 
Phytologia 21: 265. 1971. 

Crotalaria pilosa var. robusta Senn, 1939, 41: 331. Figure 
12, Map 7. 

Type: Temascaltepec, Cumbre de Tejupilco, Hinton 2686 

La Cuesta, sobre el camino a Talpa, 1,100 m, Rzedowski 15134 
(MEXU). Mexico: Plaza de Gallos, 1,200 m. Hinton 4595 (GH, NY). 
Sinaloa: Km 1206 on Mexico Hwy. 40, ca. 30 mi, E of Mazatlan- 
Guadalajara Junction, 2,700 ft., Windler & Windler 2869 (NCU). 

12. Crotalaria nayaritensis Windler, Phytologia 21: 261. 

Annual or perennial spreading herb with a taproot to 
1.5 em. thick. Stems 1-many, 6-12 dm. long, 1.5-2.5 mm. 
thick, the longest internode 3.8-10 cm. long; vestiture of 
short appressed trichomes, 0.2-0.7 mm. long. Stipules 
present, decurrent the length of the internode, 0.15-1.1 em. 
wide at the top tapering to or past the subtending node, 
the stipule lobes parallel with the stem or spreading, 0.1- 
1.3 cm. long. Leaves oval, elliptic, ovate, narrowly ovate, 
oblong or lanceolate, 3.5-7.8 cm. long, 7-26 mm. wide, base 
obtuse to cuneate, the apex obtuse, mucronate, acuminate 
or acute; vestiture of both surfaces of appressed or loosely 
appressed trichomes 0.3-0.8 mm. long; petioles 1.2-2.5 mm. 
long. Inflorescences leaf-opposed, peduncles 3.2-16 cm. long. 

1974] Crotalarias — Windler 199 

Figure 12. Crotalaria bupleurifolia var. robusta. 
A. Habit and leaf variation 
B. Stem 

200 Rhodora [Vol. 76 

Braets sessile, linear or elliptic lanceolate, 3-4.5 mm. long, 
0.2-0.5 mm. wide; pedicels 2.8-3.8 mm. long. Calyces 7.5- 
12 mm. long, the tubes 2-3.2 mm. long, bearing appressed 
trichomes 0.1-0.5 mm. long; bracteoles linear or narrowly 
lanceolate, 1.5-3 mm. long, 0.2-0.3 mm. wide, Corollas yel- 
low, the standards 7-12.5 mm. long, 2 mm. shorter to 1 mm. 
longer than the upper calyx lobes; elongate anthers 1.3-2.1 
mm. long, short anthers 0.4-0.5 mm. long; styles 5-6 mm. 
long. Fruits 1.3-2.3 em. long, 0.5-0.8 em. broad; seeds tan, 
grey-tan, green-brown, brown or red-brown, 30-35 per pod, 
1.6-2.4 mm. long. Chromosome number: n=16. Flowering 
time: August-February. Habitat: steep moist slopes and 
pine woods, elevation 2,500-6,600 feet. Range: Mexico: 
Jalisco, Nayarit. Figure 13, Map 7. 

Type: Mexico: Nayarit, North of Compostella (near 
Km. 24), about 7 miles southwest of Tepic, along road 
between Tepic and Compostella. Road-cut through moun- 
tain on moist steep slope, Windler & Windler 2902 (NCU). 

Crotalaria nayaritensis is a new species named for the 
Mexican state from which the holotype was collected. It is 
characterized by its spreading or diffuse habit, leaf-opposed 
inflorescences, small flower size, and appressed pubescence. 
Of the Mexican species it most nearly resembles C. bupleu- 
rifolia, but differs from it in having a smaller flower, 
appressed pubescence and usually narrower stipules. 

Autlan, 1,000 m, MceVaugh 19886 (MICH); Llano Verde, municipio 
de Tecalitlan, 1,600 m, Rzedowski 17417 (MiCH); 3 mi. S of Maza- 
mitla, 2,100-2,200 m, MeVaugh 12997 (micH, US); San Sebastian, 
W to Mascota, 1,425 m, Mexia 1408 (vs); Tepic, Palmer 1869 (NY, 
US). Nayarit: 10 mi. SE of Ahuacatlan, 1,100-1,200 m, Fedema. 287 
(MICH) ; N of Compostella, 3,000 ft., Windler & Windler 2902 (NCU); 
Mina Esperanza Rosa Morada, Ortega 6682 (US); 2 mi. N of Tepic, 
3,000 ft., Windler & Windler 2897 (NCU). 


During the course of the study a number of putative 
hybrids were observed, The following presentation sets 

1974] Crotalarias — Windler 201 

Figure 18. Crotalaria nayaritensis. 
A. Habit and leaf variation 
B. Stem 

202 Rhodora [Vol. 76 

forth the probable parents and lists of representative speci- 
mens which appear to be the results of crosses between 
the parents. 

Crotalaria sagittalis X C. stipularia 
ST. KITTS. Pastures near Sandy Point, Britton & Cowell 128 

Crotalaria sagittalis X C. rotundifolia var. vulgaris 
UNITED STATES. Alabama: Mobile, Mohr s.n. 1873 (SIU). 
Florida: BAY CO., E of Panama City, Windler & Windler 2723 (NCU); 
N of Betts, Windler & Windler 2721 (NCU); ESCAMBIA CO., W of 
Century, Windler & Windler 2688 (NCU); JACKSON CO., Grand Ridge, 
Windler & Windler 2712 (NCU); OKALOOSA CO. 4 mi. W of Black- 
water River Bridge on Fla. Hwy. 4, Windler & Windler 2700 (NCU); 
SANTA ROSA CO., E of Chumuckla, Windler & Windler 2693 (NCU); 
S of Milton, Windler & Windler 2697 (NcU). Mississippi: FOREST 
co., S of Hattiesburg, Windler & Windler 2659 (NCU); HARRISON CO., 
N of Biloxi, Windler & Windler 2666 (NCU); LAMAR CO., 8.5 mi. SE 
of Hattiesburg, Cooley 62 (Miss); 7.5 mi. SW of Hattiesburg, Cooper 
93 (MISS); Pearl River, 1 mi. N of Picayune, Pullen 641208 (MIss). 

Crotalaria sagittalis X C. quercetorum 
MEXICO. Chiapas: Bochil, Windler & Windler 2994 (NCU). 

Crotalaria rotundifolia X C. purshii 

UNITED STATES. Alabama: BALDWIN CO. Foley, Windler & 
Windler 2681 (NCU). Florida: CALHOUN CO., 12 mi. S of Clarksville, 
Windler & Windler 2425 (NCU); ESCAMBIA CO., 235 miles N of Jet. 
Fla. Hwy. 97 — US Hwy. 29, Windler & Windler 2686 (NCU); SANTA 
ROSA CO, S of Milton, Windler & Windler 2699 (NCU). Georgia: 
GLYNN CO., Brythe Island, Hardin & Duncan 13785 (NCU); MCINTOSH 
CO., 2.6 mi. W of Cox, Bozeman 1078 (NCU); 1 mi. SE of Ft. Bar- 
rington, Boseman & Radford 2146 (NCU); THOMAS CO, SW of 
Thomasville, Windler & Windler 2734 (NCU). Mississippi: HARRISON 
CO.: Ship Island, Demaree 35028 (LAF); JACKSON CO., Round Island, 
Demaree 33742 (LAF). 


I am especially grateful to my wife, Bonnie Kay, whose 
moral support and technical and clerical assistance speeded 
the completion of this study. 

Dr. C. Ritchie Bell, who served as research adviser during 
the study, deserves special thanks for his helpful sugges- 

1974] Crotalarias — Windler 203 

tions, criticisms, and editorial work on the manuscript. 
Drs. Albert E. Radford and Clifford R. Parks are also 
thanked for their helpful suggestions and cooperation 
during the course of the study and the preparation of the 
thesis. Dr. Velva Rudd, Smithsonian Institution, is thanked 
for her critical review of the Systematic Treatment manu- 

I would like to acknowledge my fellow students at the 
University of North Carolina at Chapel Hill and thank 
them for their aid, counsel, and criticisms during my 
years of study at the University. 

During the extensive collecting trips, help was needed 
on numerous occasions and was always cheerfully supplied 
by colleagues at other institutions. Special mention should 
be made of Dr. Arturo Gomez-Pampa, Universidad Nacional 
de México; Dr. Sidney McDaniels, Florida State University ; 
Dr. John Thieret, Southwestern Louisiana State University ; 
and Dr. D. B. Ward, University of Florida, and their 
respective staffs that provided plant dryers, and green- 
house and storage space at various times during collection 
trips. Dr. Andre Clewell and the staff of the Tall Timber 
Research Station in Tallahassee, Florida, are also due thanks 
for assistance and lodging that they provided while collec- 
tions were being made in the vicinity. 

Special thanks are extended to the curators of the 
numerous herbaria (A, ALU, AUA, BM, G, GA, GH, K, LAF, 
from which specimens were borrowed and to Dr. Norman 
K. B. Robson, in particular, whose efforts in my behalf at 
the British Museum (Natural History) in checking some 
of the early American collections proved invaluable. 


Baker, E. 1914. The African species of Crotalaria. Journal Lin- 
nean Society, Botany 42: 241-425. 

BRITTEN, J. & E. BAKER. 1897. Houstoun’s Central American Legu- 
minosae. Journal of Botany 35: 225-234. 

FERNALD, M. L. & B. SCHUBERT. 1948. Studies of American types 
in British Herbaria, IV. Some species of Thomas Walter. 
Rhodora 50: 190-208. 

204 Rhodora [Vol. 76 

GRAUSTEIN, J. 1954. The date of Pursh’s “Flora Americae Septen- 
trionalis." Rhodora 56: 275. 

LANJOUW, J. & F. STAFLEU. 1964. Index Herbariorum, Regnum 
Vegetabile, Vol. 31. 251 pp. Utrecht. 

POLHILL, R. 1968. Miscellaneous notes on African species of Cro- 
talaria L., II. Kew Bull. 22: 169-348. 

SAVAGE, S. 1945. A Catalogue of the Linnaean Herbarium. 225 
pages. London. 

SENN, H. 1939. The North American species of Crotalaria. Rhodora 
41: 317-366. 

SPRAGUE, T. 1939. Analysis of binary combinations published under 
Anonymos by Walter, Flora Caroliniana (1788). Kew Bull. 1939: 

Warp, D. 1962. The genus Anonymos and its nomenclatural sur- 
vivors. Rhodora 64: 87-92. 

WINDLER, D. R. 1970. Systematic studies in Crotalaria sagittalis 
and related species. Unpub. diss, Univ. of North Carolina at 
Chapel Hill. 258 pages. 

1971. New North American unifoliolate Crotalaria 

taxa. Phytologia 21: 257-266. 

1973. Field and garden studies in Crotalaria 

sagittalis and related species. Phytologia 26: 289-354. 




Mt. Monadnock (Elevation 3,165 feet) lies in Southern 
New Hampshire at a distance from other hills of compara- 
ble height. This mass of granite and gneiss rising from a 
plain has given rise to the generic term “monadnock” for 
similar isolated rock peaks. The Grand Monadnock, as it is 
sometimes called, to distinguish it from other monadnocks, 
is situated in the towns of Dublin and Jaffrey. Roads sur- 
round the base of the mountain at elevations of 1,200 to 
1,500 feet above sea level. 

Botanically Mt. Monadnock is of interest because of the 
presence of a few arctic-alpine plants on the bare rocky 
summit area, far from other stations for them, and because 
of the records made by Thoreau on his four visits from 
1844 to 1860, and later by other botanists, such as Stone 
(1885) and Deane (1890) and many later ones, some of 
whom deposited specimens in New England herbaria. It 
was my purpose to compare the present flora with that 
found by Thoreau. 

The student of land use can read much of its history from 
the flora, since plant habitat is affected by what has hap- 
pened to the plants. The history of Mt. Monadnock is of 
interest because of three distinct periods: (1) the develop- 
ment of complete forest cover since the time of glacial re- 
treat, (2) its complete removal on the upper slopes by fire, 
and partial removal on the lower by clearing for pasture 
and fire, and (3) the slow renewal of vegetative cover dur- 
ing the last 150 years. 

It is generally believed on the basis of geological and 
pollen evidence and observation of succession on rock and 
soil recently exposed by the retreat of present day glaciers, 
that the succession on Mt. Monadnock was probably as fol- 
lows: the first pioneer plants, following lichens and mosses, 
may have been shrubs and forbs capable of surviving under 


206 Rhodora [Vol. 76 

severe cold, wind and fluctuating moisture. These may have 
included species that colonize mineral soil and rock crevices, 
including some of the arctic-alpines. Later, as climate mod- 
erated, spruces and pines (possibly Pinus Banksiana) oc- 
cupied the accumulations of sand and organic matter in 
depressions. Finally, as humus accumulated, Red Spruce 
(Picea rubens) came to dominate the flora of the upper part 
of the mountain where soil development was indefinitely 

Mt. Monadnock was completely forested at the coming of 
the White Man, the summit area covered with Red Spruce 
to the very summit, and the lower slopes with spruce and 
hardwood, much as the case of lesser mountains much 
farther north. All similar rocky peaks below the approxi- 
mately 4,500 feet climatic timberline in New Hampshire 
are or were completely forest-clad. Mt. Monadnock is about 
1,000 feet lower than climatic timberline and lies almost 
100 miles south of the nearest timberline. Tree line is 
higher at lower latitudes. The bare rocky summits of New 
Hampshire such as Chocorua, Cardigan, Firescrew, and 
Percy Peaks all have documented history of fire. 

Until recent years there were many old stumps and logs 
near the summit. Thoreau wrote in his Journal (June 
1858) : “The whole rocky part of the mountain, except the 
extreme summit, is strewn with the relics of spruce trees a 
dozen or fifteen feet long, and long since dead and bleached, 
so that there is plenty of fuel at hand." The remains near- 
est the summit were no doubt first consumed by visitors 
campfires. Visitors were very numerous in Thoreau’s day 
and before, and in several of his writings he reports having 
seen fires at the top. Since the area at the extreme summit 
is quite level, with depressions, it is probable that it was 
originally as densely forested as the rest of the mountain. 
Chamberlain (1936) shows a picture (p. 73) of stumps near 
the top. Obviously, there could not have been such severe 
fires at the summit had there not been timber to burn. The 
1953 fire on the east slope stopped when it reached the 
scrub zone where there was no fuel. 

1974] Flora of Mount Monadnock — Baldwin 207 

There are many published reports that confirm the orig- 
inal forested nature of the summit. Later the practice of 
clearing the slopes for pasture gave rise to numerous fires. 

During the celebration of the centennial of the Town of 
Dublin in 1852, Charles Mason made an address on June 17 
(Leonard 1855) including the following remark: 

“Formerly covered with trees almost to the summit.” 

At the same celebration in Dublin (Leonard op. cit.) a 
letter was read from Daniel Elliott, M.D. in which he re- 
called: “I remember (Monadnock) when clothed with ver- 
dant foliage to the very summit.” Similar records are 
found in the Town of Jaffrey: “When the town was first 
settled the mountain was covered to its summit with forest 
trees, principally spruce, excepting a small peak southeast 
of the top, which was called the “Bald Rocks” (Cutter 
1881). “Belts of heavy spruce timber on the highest por- 
tions” were reported by Child (1885). Similar references 
may be found in Dana (1816) and Ellis (1880). 

Ecologists recognized that forest had come to occupy the 
mountain much as peaks farther north. Braun (1950) 
refers to Monadnock (p. 423) as “covered with Red Spruce 
from 2,300 feet upward, thus producing a small outlier of 
the prevailing mountain vegetation in the north.” Spurr 
(1964) states that “the summit of Mt. Monadnock ... was 
clothed with a Red Spruce forest until destroyed by fire...” 

Many fires have occurred during the recorded history of 
Mt. Monadnock, and it is possible that lightning fires may 
have burned in earlier times, but it is doubtful if Indians 
caused any fires on its forbidding slopes. In the same ad- 
dress referred to above (Leonard, loc. cit.) Charles Mason 
said: “A great fire spread over a great part of the superior 
portion of the mountain killing every tree and shrub. Dead 
trees falling, furnished materials for another conflagration. 
Some 30 years ago in the latter part of a dry summer, a fire 
in a clearing made its way up to the higher regions." “Some 
30 years ago” would have placed this fire about 1820, al- 
though the cause of the fire differs from other accounts 
given below. Also the statement of Dr. Elliott on the same 

208 Rhodora [Vol. 76 

occasion: “I saw, year after year, the devouring flames 
climbing his lofty sides", suggests many fires. 

Thoreau, on his second visit to Monadnock in 1852 (his 
first was in 1844, the journal report of which has not been 
preserved) related this incident during his passage through 
Peterborough: *A man in Peterborough told me that his 
father told him that Monadnock used to be covered with 
forest; that fires ran through it and killed the turf; then 
the trees were blown down, and their roots turned up and 
formed a dense and impenetrable thicket in which wolves 
abounded . . . till finally they set fire to this thicket and it 
made the greatest fire they had ever had in the county.” 

These and other records fail to give exact dates. As 
Chamberlain (1936) says (p. 97): “No documentary evi- 
dence has been found to show exactly when the fires oc- 
curred.” He also quotes (p. 98) from the New Hampshire 
Sentinel of September 30, 1825, reporting a great fire of 
“some years ago”. “This fire consumed only the turf. The 
trees turned over in every direction, and afforded about 
three years later fuel for a more tremendous scene.” This 
places the two “great” fires closer together. The best evi- 
dence suggests a first fire about 1800, then the 1815 hurri- 
cane, and the final great fire in 1820, There were later ones, 
too. Thoreau reported (“A Walk to Wachusett”, July 19, 
1842) “A fire blazing on Monadnock that night, which 
lighted up the whole western horizon.” 

The first inroads of man occurred on the lower slopes that 
were cleared for pasture. Later, clearing progressed higher 
where sheep were grazed. Fire was used both for opening 
the areas and for keeping them so, and many of the fires 
that reached the upper levels both before and after the 
1820 fire were caused by fire escaping from pasture burn- 
ing. The repeated burning promoted the growth of blue- 
berries, which I believe may have spread from lower 
pastures to the summit after the great fires. At present 
these are rather scarce. 

Following abandonment of the pastures, these seeded into 
Red Spruce, and the process of reclothing the upper reaches 

1974] Flora of Mount Monadnock — Baldwin 209 

of the mountain progressed rapidly at first, and ever slower 
on the more exposed parts of the mountain. Here denuda- 
tion resulted in a worsening of the climate. Wind movement 
increased over the bare rock, and extremes of temperature 
and soil moisture retarded growth. Recently a cycle of 
retarded spread of spruce seems to have set in, since almost 
no spruce trees less than forty years old can now be found 
on the upper part of the mountain and a high percentage 
of the trees display dead tops and reduction in diameter 
and height growth. 

The large area laid bare by the fires apparently provided 
a favorable habitat for a few arctic-alpine plants that may 
have been present before the fires or had been transported 
from other sources. In any case they were well-established 
at the time of Thoreau's first visit, little over twenty years 
after the great fire. Those recorded by Thoreau (Journal, 
August 8, 1860, 25-28; 1670-1671) were: Arenaria 
groenlandica, Vaccinium Vitis-Idaea var. minus and Juncus 
trifidus. Others with northern or high altitude affinities, 
not found generally in the lowland, especially in the vicinity 
of Mt. Monadnock were Potentilla tridentata and Smilacina 
trifolia. During the present survey Lycopodium Selago var. 
Selago and var. appressum as well as L. annotinum var. 
pungens have been added to the list. 

Thoreau limited his observations to the "upper part of 
the mountain," and essentially all the taxa reported by him 
in 1858 “within 200 feet of the summit" were also collected 
in 1968. Less complete were the lists of Stone (1885), 
Deane (1890) and Emerson (1869), probably due to the 
restricted area that they examined. While almost all the 
taxa recorded earlier are present now (at least those easily 
discoverable), and a substantial number of new ones have 
been added, there are doubtless more to be found. There 
have been changes in distribution and abundance of the 
plants recorded by the early botanists. The encroachment 
of Red Spruce and more especially shrubs such as Nemo- 
panthus mucronata, Rhododendron canadense and Kalmia 
angustifolia appears to have reduced the habitat available 

210 Rhodora [Vol. 76 

to the arctic-alpines. Vaccinium Vitis-Idaea can persist a 
long time under these shrubs, but fruiting is diminished. 
Some thirty years ago the proprietors of the Half Way 
House used to collect enough mountain cranberries on the 
mountain to supply hotel guests, while now only small 
quantities can be picked. This plant has almost disappeared 
from the southwest side where the hotel supply was largely 
obtained. The alpine flora seems to be increasingly re- 
stricted to the northern side of the peak where a cooler 
climate prevails. Further, observations indicate that there 
may be a successional change going on in the mats of Vac- 
cinium Vitis-Idaea, that are invaded more and more by 
Potentilla tridentata. Another major change from 100 
years ago is the vast reduction in Cladonia lichens. Thoreau 
remarks frequently how dense they were, crunching under 
foot, and dry so that fire would spread through them. 
Cladonia persists around the margins of shrubby hollows, 
but today does not cover extensive areas. Other lichens 
mentioned by Thoreau on the summit are now mostly con- 
fined to moist ravines. 

Originally it was planned to limit collections to the area 
studied by Thoreau, “the upper part of the mountain". Be- 
cause of the difficulty of determining just what this em- 
braced, the area was extended to include land above 2,000 
feet above sea level. (The Half Way House site is at 2,100 
feet) Present collections have been supplemented by recent 
herbarium records kindly supplied by Mr. and Mrs. David 
E. Boufford of Keene State College. Where several collec- 
tions of the same taxon were found, only the earliest record 
is given, in order to save space.* All records that stated 
Mt. Monadnock as the place of collection were included, 
although often the specific location was not given. Genera 
and species are listed in the order of Gray's Manual, 8th 
Edition (Fernald 1950). The following symbols are used: 

(*) asterisk: reported by Thoreau 1858-1860. 

(—) dash: collected 1968-1972. 

*Complete lists have been deposited in several New England her- 

1974] Flora of Mount Monadnock — Baldwin 211 

(+) dagger: collected 1968-1972, not previously reported 
(as far as known) 

Most of the specimens collected in 1968-1972 were de- 
posited in the University of New Hampshire and Franklin 
Pierce College herbaria. Abbreviations for all Herbaria 
consulted in this study are as follows: HNH Dartmouth 
College, FPC Franklin Pierce College, KSC Keene State Col- 
lege, NEBC New England Botanical Club, MASS University 
of Massachusetts, and NHA University of New Hampshire. 


-Lycopodium Selago L. var. Selago L. W. Deane, Aug., 
1891 (NEBC). 
+_L. Selago L. var. appressum Desv. H. I. Baldwin, July. 
1968 (FPC). 
* L.lucidulum Michx. H. I. Baldwin, Aug., 1968 (FPC, 
+—L. inundatum L. H. I. Baldwin, Oct., 1968 (FPC). 
* L.annotinum L. H. D. Thoreau, Aug., 1869 (?) (NEBC). 
+_L. annotinum L. var. pungens (La Pylaie) Desv. H. I. 
Baldwin, June, 1971 (FPC, NHA). 
*_L. clavatum L. T. L. Steiger, Oct., 1935 (NHA). 
L. clavatum L. var. megastachyon Fern. & Bissell, C. 
Jenks, Aug. 1883 (NEBC). 
_L. obscurum L. var. obscurum. A. W. Driggs. Aug., 1916 
L. obscurum L. var. dendroideum (Michx.) D. C. Eat. 
B. L. Robinson, Aug., 1869 (NEBC). 
*_L. complanatum L. A. W. Driggs, Aug., 1916 (NEBC). 
Botrychium matricariaefolium A. Br. E. L. Rand, May, 
1897 (NEBC). 
i-Osmunda cinnamomea L. H. I. Baldwin, Aug., 1968 (FPC). 
Woodsia ilvensis (L.) R. Br. B. L. Robinson, Aug., 1898 
—Thelypteris Phegopteris (L.) Slosson. E. F. Williams, 
June, 1894 (NEBC). 
—Dryopteris spinulosa (O. F. Muell.) Watt. var. americana 
(Fisch.) Fernald. E. F. Williams, June, 1894 (NEBC). 

212 Rhodora [Vol. 76 

—D. marginalis (L.) Gray. D. S. Boufford, June, 1971 (KSC). 
+—Polystichum acrostichoides (Michx.) Schott. H. I. Bald- 
win, Aug., 1971 (FPC, NHA). 
*—Dennstaedtia punctilobula (Michx.) Moore. H. I. Baldwin, 
Oct., 1968 (FPC). 

-Pteridium aquilinum ( L.) Kuhn var. latiusculum (Desv.) 
Underwood. H. I. Baldwin, Aug., 1968 (FPC). 

*—Polypodium virginianum L. W. Deane, Aug., 1889 (HNH). 

"—Abies balsamea (L.) Mill, J. R. Churchill, May, 1895 

*—Tsuga canadensis (L.) Carr. H. I. Baldwin, July, 1968 

*—Picea rubens Sarg. E. F. Williams, May, 1898 (HNH). 

T-Larix laricina (DuRoi) K. Koch. H. I. Baldwin, July, 
1968 (FPC). (one tree at 2,900 feet on west side, two 
in Thoreau Bog). 

*-Pinus Strobus L. A. R. Hodgdon et al., Aug., 1956 (NHA). 

Poa compressa L. W. Deane, July, 1884 (HNH). 
P. pratensis L. E. F. Williams, June, 1894 (NEBC). 
P. palustris L. J. R. Churchill, July, 1890 (MASS, NEBC). 

*—_Festuca ovina L. E. L. Dickinson and F. C. Seymour, 

July, 1930 (Mass). 
T-F. rubra L. H. I. Baldwin, July, 1968 (FPC). 
*_Deschampsia flexuosa (L.) Trin. H. D. Thoreau, Aug., 
1860 (NEBC). 
*_Danthonia spicata (L.) Beauv. H. D. Thoreau, Aug., 1860 
—D. compressa Aust. A. S. Goodale, July, 1927 (MASS). 
*Calamagrostis canadensis ( Michx.) Nutt. H. D. Thoreau, 
Aug., 1860 (NEBC). 

*—Agrostis scabra Willd. H. D. Thoreau, Aug., 1860 (NEBC). 
-Scirpus atrovirens Willd. C. W. J enks, Aug., 1883 (NEBC). 
—S. atrocinctus Fern. A. S. Goodale, July, 1927 (Mass). 

*—Eriophorum spissum Fern., H. D. Thoreau, Aug., 1860 

(?) (NEBC). 
*E. gracile W. D. J. Koch. A. S. Goodale, July, 1927 (MASS). 
-E. tenellum Nutt. R. M. Harper, July, 1899 (NEBC). 
“3,150 feet”. 

1974] Flora of Mount Monadnock — Baldwin 213 

E. viridi-carinatum (Engelm.) Fern. C. F. Batchelder, 
Sept., 1909 (NHA). 
—E. virginicum L. C. W. Jenks and C. W. Swan, Aug., 
1883 (NEBC, NHA). 
Rhynchospora capitellata (Michx.) Vahl. B. L. Robinson, 
July, 1897 (NEBC). 
+—R. alba (L.) Vahl. H. I. Baldwin, Oct., 1968 (FPC). 
+—Carex stipata Muhl. H. I. Baldwin, June, 1969 (FPC). 
*C. trisperma Dew. H. D. Thoreau, Aug., 1860 (NEBC). 
“NE swamp”. 
+-C. trisperma Dew. var. Billingsii Knight, H. I. Baldwin, 
Oct., 1968 (FPC). 
* C. canescens L. “E. F. Jr." (?), 1838 (MASS). 
C. canescens L. var. subloliacea Laested. Sydney Harris, 
June, 1894 (NEBC). 
C. brunnescens (Pers.) Poir. W. Deane, July, 1889 (MASS). 
C. brunnescens (Pers.) Poir. var. sphaerostachya (Tuk- 
erm.) Kükenth. A. W. Driggs, Aug., 1916 (NEBC). 
C. cumulata (Bailey) Mackenz. C. W. Jenks, Aug., 1885 
—C. communis Bailey. L. Russell, 1898 (NHA). 
C. novae-angliae Schwein. W. Deane, July, 1891 (HNH, 
MASS). “Mt. road below house". 
C. deflexa Hornem. J. R. Churchill, May, 1895 (NEBC). 
C. umbellata Schkuhr. J. R. Churchill, May, 1896 (NEBC). 
C. gracillima Schwein. R. J. Eaton, July, 1957 (NHA). 
C.aestivalis M. A. Curtis. J. R. Churchill, July, 1890 
C. leptonervia Fern. E. F. Williams, June, 1894 (NEBC). 
C. intumescens Rudge. E. F. Williams, June, 1894 (NEBC). 
+-C. nigra L. H. I. Baldwin. July, 1971 (FPC, NHA). 
i-Arisaema triphyllum (L.) Schott. D. W. Allen, May, 1968 
-Juncus bufonius L. A. W. Driggs, Aug., 1916 (NEBC). 
*_J. trifidus L. C. W. Swan, Aug., 1883 (NEBC). 
J.Greenei Oakes and Tuckerm. B. L. Robinson, Aug., 
1896 (NEBC). 

214 Rhodora [Vol. 76 

-J. brevicaudatus (Engelm.) Fern. A. W. Driggs, Aug., 

1916 (NEBC). 
J. articulatus L. C. A. and A. W. Cheever, Sept., 1910 

( NEBC). 

T-Luzula, multiflora (Retz.) Lejeune. D. W. Allen, May, 
1968 (FPC, NHA). 

*—Uvularia sessilifolia L. D. W. Allen, May, 1968 (FPC). 

*—Clintonia borealis (Ait.) Raf. T. L. Steiger, May, 1937 

r-Smilacina racemosa (L.) Desf. H. I. Baldwin, Sept., 1970 

*—S. trifolia (L.) Desf. Sydney Harris, June, 1894 (NEBC). 

r-Maianthemwm canadense Desf. D. W. Allen, May, 1968 

—Streptopus roseus Michx. var. perspectus Fassett. W. 
Deane, July, 1889 (NEBC). 

*-Polygonatum pubescens (Willd.) Pursh. H. I. Baldwin, 
May, 1969 (FPC, NHA). 

-Medeola virginiana L. D. W. Allen, May, 1968 (FPC). 

-Trillium erectum L. J. R. Churchill, May, 1883 (Mass). 

*—T. undulatum Willd. T. L. Steiger, May, 1937 (NHA). 

r-Sisyrinchium montanum Greene. H. I. Baldwin, July, 

1968 (NHA). 
*-Cypripedium acaule Ait. D. W. Allen, May, 1968 (FPC, 
Habenaria Hookeri Torr. Henry A. Purdie, June, 1905 

Spiranthes lacera Raf. Hodgdon and Ken Welsh, July, 
1957 (NHA). “2,500 feet”. 
Malaxis uniflora Michx. F. Hunt, Aug., 1946 (NHA). 
*Salix Bebbiana Sarg. A. R. Hodgdon, June, 1933 (NHA). 
*—S. discolor Muhl. D. W. Allen, May, 1968 (FPC). 
*—Populus tremuloides Michx. H. I. Baldwin, Aug., 1968 
*_Betula alleghaniensis Britton. Hodgdon and Welsh, July, 
1957 (NHA). 
-B. populifolia Marsh. Hodgdon et al., Aug., 1956 (NHA). 
*—B. papyrifera Marsh. Hattie Merrifield, July, 1879 (NEBC). 

1974] Flora of Mount Monadnock — Baldwin 215 

—B. papyrifera Marsh. var. cordifolia (Regel) Fern. Hodg- 
don and Welsh, July, 1957 (NHA). 
B. papyrifera Marsh. X B. populifolia Marsh. Hodgdon 
et al., Aug., 1956 (NHA). “2,800 feet". 
*_Ostrya virginiana (Mill. K. Koch. H. I. Baldwin, Aug., 
1971 (FPC). 
—Fagus grandifolia Ehrh. Hodgdon et al., Aug., 1956 (NHA, 
i-Quercus rubra L. D. W. Allen, May, 1968 (FPC). 
Polygonum aviculare L. var. angustissimum Meisn. B. L. 
Robinson, July, 1897 (NEBC). 
* P. cilinode Michx. W. Deane, Aug., 1889 (HNH). 
+_P. scandens L. H. I. Baldwin, Aug., 1972 (FPC, NHA). 
“one plant, Red Spot Trail”. 
+_Claytonia caroliniana Michx. H. I. Baldwin, June, 1971 
*_Arenaria groenlandica Du Roi. Thomas Hope, Aug., 1879 
Ranunculus abortivus L. Rand and Robinson, June, 1898 
+—R. acris L. H. I. Baldwin, July, 1971 (NHA). 
+_R. bulbosus L. D. W. Allen, June, 1968 (NHA). 
i-Thalictrum polygamum Muhl. H. I. Baldwin, May, 1969 
*_Coptis groenlandica (Oeder) Fern. D. W. Allen, May, 
1968 (FPC). 
3—Actaea pachypoda Ell. H. I. Baldwin, May, 1969 (FPC). 
*_Aquilegia canadensis L. Rand and Robinson, May, 1897 


-Berberis Thunbergii DC. W. H. Manning, Oct., 1913 

_Drosera intermedia Hayne. Fred M. Hunt, Aug., 1946 

+_D. rotundifolia L. H. I. Baldwin, July, 1968 (FPC, NHA). 
+-Chrysosplenium americanum Schwein. D. E. Boufford, 
1971 (KSC). 
*Ribes cynosbati L. T. Otis Fuller, June, 1883 (NEBC). 
* R. glandulosum Grauer. J. R. Churchill, May, 1883 (NHA). 

216 Rhodora [Vol. 76 

R. triste Pall. T. L. Steiger, May, 1937 (NHA). 
*_Spiraea latifolia (Ait.) Borkh. A. W. Driggs, Aug., 1916 
(NEBC). "above timberline". 
1-8. tomentosa L. H. I. Baldwin, July, 1968 (FPC). 
*—Pyrus melanocarpa (Michx.) Willd. C. F. Batchelder. 
Sept., 1909 (NHA). 
*—P. americana (Marsh.) DC. J. R. Churchill, July, 1890 
P. decora, (Sarg.) Hyland. Hodgdon and Kenneth Welsh, 
July, 1957 (NEBC). ‘2,500 feet". 
P. americana L. X P. decora (Sarg.) Hyland. Hodgdon 
and Kenneth Welsh, July, 1957 (NEBC). “2,500 feet". 
*—Amelanchier laevis Wieg. Rand and Robinson, May, 1897 
*_A. Bartramiana (Tausch.) Roemer. J. R. Churchill, May, 
1885 (NEBC). 
A. Bartramiana (Tausch.) Roemer. X A. neglecta (?) 
Rand and Robinson, May, 1897 (NEBC). 
T-Crataegus macrosperma Ashe. D. W. Allen, May, 1968 
T-Fragaria virginiana Duchesne. D. W. Allen, May, 1968 
Potentilla fruticosa L. J. A. Bates, Aug., 1900 (NEBC). 
*—P. tridentata Ait. E. S. Hoar, July, 1857 (NEBC). 
P. norwegica L. A. W. Driggs, Aug., 1916 (NEBC). “sum- 
T—P. canadensis L. H. I. Baldwin, June, 1969 (FPC, NHA). 
T—P. simplex Michx. H. I. Baldwin, May, 1970 (FPC). 
Geum aleppicum Jacq. var. strictum (Ait.) Fern. Henry 
Purdie, July, 1905 (HNH). 
Rubus hispidus L. J. P. Poole, July, 1939 (HNH, NHA). 
R. setosus Bigel. A. W. Driggs, Aug., 1916 (NEBC). 
*_R. vermontanus Blanch. B. L. Robinson, July, 1897 (NEBC). 
R. allegheniensis Porter. A. W. Driggs, Aug., 1916 (NEBC). 
T—R. frondosus Bigel. H. I. Baldwin, July, 1968 (FPC). 
*—Prunus pensylvanica L. f. C. W. Jenks, Aug., 1883 (NEBC). 
T—P. serotina Ehrh. H. I. Baldwin, July, 1968 (FPC). 
t-Trifolium repens L. D. W. Allen, June, 1968 (FPC). 

1974] Flora of Mount Monadnock — Baldwin 217 

_Oxalis montana Raf. T. Otis Fuller, June, 1883 (NHA). 
Polygala sanguinea L. R. H. Piper, July, 1927 (MASS). 
* Nemopanthus mucronata (L.) Trel. Rand and. Robinson, 
May, 1897 (NEBC). 
* Acer spicatum Lam. Sydney Harris, June, 1894 (NEBC). 
* A. pensylvanicum L. C. W. Jenks, Aug., 1883 (NEBC). 
1—A. saccharum Marsh. H. I. Baldwin, June, 1968 (FPC). 
* A. rubrum L. H. I. Baldwin, June, 1968 (FPC, NHA). 
Hypericum perforatum L. J. P. Poole, July, 1939 (HNH). 
_H. canadense L. Mary E. Gilbreth, Aug., 1889 (NEBC). 
(one extremely small plant found in 1971, H. I. Bald- 
+—Viola papilionacea Pursh. H. I. Baldwin, May, 1969 (NHA). 
V. sororia Willd. E. F. Williams, May, 1898 (NEBC). 
V. fimbriatula Sm. E. F. Williams, May, 1898 (NEBC). 
—V. pallens (Banks) Brainerd. A. W. Cheever, Sept., 1910 
V. blanda Willd. J. R. Churchill, July, 1890 (HNH, NEBC). 
V. lanceolata L. A. R. Hodgdon, June, 1933 (NHA). 

1-V. rotundifolia Michx. H. I. Baldwin, April, 1969 (NHA). 
V. adunca Sm. Rand and. Robinson, May, 1897 (NEBC). 
1—V. conspersa, Reichenb. H. I. Baldwin, May, 1971 (FPC). 

Circaea alpina L. Edward S. Hoar, 1879 (NEBC). 
* Epilobium angustifolium L. H. I. Baldwin, Aug., 1971 
E. leptophyllum Raf. C. W. Jenks, Aug., 1883 (NEBC). 
Aralia racemosa L. Hattie Merrifield, July, 1879 (NEBC). 
* A4. hispida Vent. J. P. Poole, July, 1939 (HNH, NHA). 
_A. nudicaulis L. A. S. Goodale, May, 1926 (MASS). 
1—Panaz trifolius L. D. W. Allen, May, 1968 (FPC, NHA). 
* Cornus canadensis L. C. W. Jenks, Aug., 1883 (NEBC). 
—C. alternifolia L. f. Hodgdon et al., Aug., 1956 (NHA). 
Chimaphila umbellata (L.) Bart. var. cisatlantica Blake. 
J. P. Poole, July, 1939, (HNH, NHA). 
+-Monotropa uniflora L. H. I. Baldwin, Sept., 1968 (FPC). 
* Ledum groenlandicum Oeder. Sydney Harris, June, 1894 

218 Rhodora [Vol. 76 

*—Rhododendron canadense (L.) Torr. Sydney Harris, June, 
1883 (NEBC). 
*—-Kalmia angustifolia L. J. P. Poole, July, 1939 (HNH). 
*-Cassandra calyculata (L.) D. Don. D. W. Allen, May, 
1968 (FPC). 
*—Epigaea repens L. H. I. Baldwin, April, 1969 (FPC). 
“(one plant only, on cliff) ." 
*Gaultheria hispidula (L.) Bigel. C. G. Trow, 1871 (MASS). 
1T-Calluna vulgaris (L.) Hull. H. I. Baldwin, May, 1965 
(NHA). (one isolated colony (introd)). 
-Gaylussacia baccata (Wand.) K. Koch. A. W. Driggs, 
Aug., 1916 (NEBC). 
Vaccinium myrtilloides Michx. Hodgdon et al., Aug., 1956 
(NHA). “2,800 feet", 
*_V. angustifolium Ait. W. Deane, Aug., 1889 (HNH). 
-V. angustifolium Ait. var. laevifolium House. B. L. Rob- 
inson, July, 1897 (NEBC). 
T-V. augustifolium Ait, X V. corymbosum L. H. I. Baldwin, 
Aug., 1969 (NHA). 
V. angustifolium Ait. var. nigrum ( Wood) Dole. Fred M. 
Hunt, Aug., 1946 (NHA). 
V. corymbosum L. A. S. Goodale, May, 1926 (Mass). 
*-V. Vitis-Idaea L. var. minus Lodd. C. G. Trow, Aug., 1864 
T—Asclepias syriaca L. H. I. Baldwin, July, 1971 (FPC). 
*—Trientalis borealis Raf. D. W. Allen, May, 1968 (FPC). 
Lysimachia terrestris (L.) BSP. J. P. Poole, July, 1939 
(HNH, NHA). 
T-Fraxinus americana L. H. I. Baldwin, July, 1968 (FPC). 
Hedeoma pulegioides (L.) Pers. B. L. Robinson, Aug., 
1896 (NEBC). 
f-Veronica longifolia L. Hodgdon and Hehre, Aug., 1969 
1—-V. officinalis L. H. I. Baldwin, June, 1969 (FPC, NHA). 
Gerardia tenuifolia Vahl. B. L. Robinson, Aug., 1898 
*“_Melampyrum lineare Desr. var. americanum (Michx.) 
Beauv. D. W. Allen, June, 1968 (FPC). 

1974] Flora of Mount Monadnock — Baldwin 219 

1-Mitchella repens L. H. I. Baldwin, Aug., 1968 (FPC, NHA). 
tT-Houstonia caerulea L. D. W. Allen, May, 1968 (FPC, NHA). 
*_Diervilla Lonicera Mill. C. W. Jenks, Aug., 1883 (NEBC). 
*_Viburnum alnifolium Marsh. C. W. Jenks, Aug., 1883 
*_V. cassinoides L. C. L. Canfield, June, 1931 (NHA). 
*_Sambucus pubens Michx. W. Deane, July, 1891 (NEBC). 
T-Solidago caesia L. H. I. Baldwin, Sept., 1970 (FPC). 
*_S. macrophylla Pursh. W. Deane, Aug. 1891 (HNH, NEHA, 
—S. puberula Nutt. Mary E. Gilbreth, Aug., 1889 (NEBC). 
—S. Randii (Porter) Britt. C. W. Jenks, Aug., 1883 (NHA). 
S. Randii (Porter) Britt. var. monticola Porter. C. F. 
Batchelder, June, 1919 (NHA). 
S. juncea Ait. Parsons and. Seymour, Aug., 1927 (MASS). 
—S. nemoralis Ait. Fred M. Hunt, Aug., 1945 (NHA). 
*—S. rugosa Ait. var. rugosa H. I. Baldwin, Oct., 1968 (NHA). 
*_S. graminifolia (L.) Salisb. H. I. Baldwin, Sept., 1968 
(FPC, NHA). 
T—Aster divaricatus L. H. I. Baldwin, Sept., 1968 (FPC). 
*—A. acuminatus Michx. B. L. Robinson, Aug., 1896 (NEBC). 
T-Erigeron pulchellus Michx. H. I. Baldwin, July, 1968 

*—Anaphalis margaritacea (L.) B.&H. D. W. Allen, May, 
1968 (FPC). 
Gnaphalium Macounii Greene. C. W. Jenks, Aug., 1883 

G. obtusifolium L. C. A. Cheever, Sept., 1910 (HNH). 

G. uliginosum L. W. Deane, Aug. 1891 (HNH). 
*—Achillea Millefolium L. D. W. Allen, May, 1968 (FPC). 
t-Chrysanthemum Leucanthemum L. var. pinnatifidum Le- 

coq and Lamotte. H. I. Baldwin, July, 1971 (FPC, NHA). 

Taraxacum erythrospermum Andrz. E. F. Williams, May, 

1898 (HNH). 
—T. officinale Weber. E. F. Williams, May, 1898 (NEBC). 
T-Prenanthes trifoliata (Cass.) Fern. var. nana (Bigel.) 
Fern. D. W. Allen, May, 1968 (FPC). 
1—P. altissima L. H. I. Baldwin, Aug., 1968 (NHA). 



[Vol. 76 

Additional taxa reported by Thoreau in his journals of 
1858 and 1860. The names, as given by Thoreau, including 
his question marks are at the left in the following list: 
Contemporary accepted binomials are listed at the right 
opposite the name he used if that name is no longer used 
or if only a common name were given by him. 

Sensitive fern 
Black spruce 

Cinna arundinacea 
Glyceria elongata 

Eleocharis tenuis 
Scirpus Eriophorum 
Carex straminea 

C. scoparia ? 

C. debilis 

C. Deweyana 
Juncus acuminatus ? 

J. parodoxus ? 

Salix lucida 
S. humilis 
Tiarella cordifolia 
Amelanchier canadensis 
var Botryapium 
A. canadensis 
var oblongifolia 
Geranium Robertianum 
Epilobium palustre 
Water andromeda 

Chelone glabra 
Solidago virgaurea 

Onoclea sensibilis L. 

No Picea mariana (Mill) BSP. 
has been found since Tho- 
reau’s visits; this may be P. 
rubens Sarg. 

G. melicaria (Michx.) F. T. 


S. cyperinus (L.) Kunth. 
C. tenera Desv. 

J. acuminatus Michx. or J. can- 
adensis J. Gay 

perhaps J. canadensis or J. acu- 

A. arborea (Michx. F.) Fern. 

A. canadensis (L.) Medic. 

A. glaucophylla Link or Kalmia 
polifolia Wang. 

Gaultheria procumbens L. 


1974] Flora of Mount Monadnock — Baldwin 221 

Three-ribbed goldenrod 

(not canadensis) S. altissima L. or S. gigantea Ait. 
Aster thyrsoidea ? 
Diplopappus umbellatus Aster umbellatus Mill. 


The present flora of the upper part of Mt. Monadnock is 
as noteworthy for the absence of certain plants as by the 
presence of unusual ones such as the arctic-alpines. Thus 
Alnus has not been found anywhere on the slopes of the 
mountain, nor has Picea mariana been recorded from the 
spruce-lined bogs. Acer saccharum is confined to a small 
area on the east side. Further there are many instances of 
lowland adventives that take root and may thrive for a 
time and then disappear. This may explain why some spe- 
cies recorded earlier have not now been found (aside from 
less keen observation). A total of 220 taxa are represented 
by specimens in herbaria, including 152 taxa collected from 
1965 to 1972. Fifty-nine taxa collected during this period 
were not previously found in herbaria, and may possibly 
have been collected for the first time on the mountain. Of 
the 94 taxa reported by Thoreau, 68 of which are in her- 
baria, 62 were collected during the present survey. 

Thoreau observed seeds of many lowland plants blowing 
over the summit, and reflected on the origin of adventives 
(and perhaps transitory taxa) : 

“I saw what I took to be a thistle-down going low over 
the summit, and might have caught it, though I saw no 
thistle on the mountain top, nor any other plant from which 
this could have come. (I have no doubt it was a thistle by 
its appearance and its season.) It had evidently come up 
from the country below. This shows that it may carry its 
seeds to higher regions than it inhabits, and it suggests 
how the seeds of some mountain plants, such as the Solidago 
thyrsoidea may be conveyed from mountain to mountain, 
also other solidagos, asters, epilobiums, willows, etc." Tho- 
reau: Journal August 9, 1860, (p. 50) (page 1677 in the 
1962 Dover edition). 

222 Rhodora [Vol. 76 

It seems obvious that most of the wind-disseminated spe- 
cies that have restocked the mountain since the 1820 fire, 
including Red Spruce, have come by the aerial route, mostly 
probably from the adjacent lowland. Light-seeded arctic- 
alpines may be included. It is less easy to explain how 
heavy-seeded plants such as Vaccinium arrived. These may 
have been present before the fires, surviving the shade of 
spruce on some steep ledges, or they may have been carried 
by birds migrating from timberline areas in the White 

According to Pease (1964 p. 15) the following taxa col- 
lected or reported from Mt. Monadnock “may be listed as 
essentially alpine": 

Lycopodium Selago (including var. appressum) 

Lycopodium annotinum var. pungens 

Juncus trifidus 

Arenaria groenlandica 

Solidago macrophylla var. thyrsoidea 
Mt. Monadnock appears to be the southern limit for some 
of these. 

The relation of fire to the occurrence of arctic-alpine 
plants needs more investigation. It is curious that they be- 
came so abundant on Mt. Monadnock 29-30 years after the 
great fire. Do the ashes from fire favor their establishment 
(Fernald (1907) thought alkaline soil most unfavorable) 
or is it open space, exposed rock crevices, or mineral soil 
alone? Many years ago, searching for Vaccinium Vitis- 
Idaea on Cape Breton, I could find it only on an area that 
had been swept by fire about ten years before. 

There are thus at least two possible explanations for the 
presence on Mt. Monadnock of taxa usually found above 

(1) They have survived since post-glacial times in re- 
fugia on the mountain itself possibly on lower slopes as pro- 
posed by Fernald (1907, 1925), and then survived the 
fires (which were very severe indeed), and then restocked 
the bare rock and open soil. 

(2) The summit flora may have arisen from disseminules 

1974] Flora of Mount Monadnock — Baldwin 223 

carried from timberline areas by wind, animals or birds, 
or even men. 

Steele and Hodgdon (1973) reached the same possible 
explanations for the presence of Carex capitata and Carex 
Bigelowii on Mt. Cardigan. 

Regardless of the origin of the present flora, processes 
of change are at work, reducing the abundance and area 
occupied by some plants, while others are assuming dom- 
inance. Many lowland forms may invade and thrive for a 
time and then disappear. Neither the rate nor direction of 
succession is uniform. Invasion of spruce on the upper 
rocks has slowed to a temporary halt, even while the extent 
of arctic-alpine plants declines. 

The Problem of the Source and Persistence of 
Arctic-Alpine Plants 

How did arctic-alpine plants survive from post-glacial 
times on lower mountains and escape competition with the 

One explanation for the presence of arctic-alpine plants 
below timberline is that they survived in refugia, rock 
crevices or smal] exposed ledges since post-glacial times, 
and that these crevices and ledges did not become over- 
grown by other plants and that the microclimate remained 
favorable. Then, when more open habitat was exposed by 
fire, these plants re-invaded the bare summits and increased 
greatly. Fernald (1907) postulated that alpine plants, 
“forced to find temporary homes during the ice age on 
potassic, calcareous or magnesium areas, found in the allu- 
vial deposits a mixed soil in which they were able to spread. 
As climate at low altitudes became warmer, these plants 
found on cold cliffs and exposed mountain summits the only 
habitats in which they have been able to persist within the 
temperate regions” It is fairly easy to understand that they 
were able to persist on alpine tundra above timberline, and 
indeed on some “cold cliffs’ like those in Dixville Notch, 
New Hampshire, and possibly elsewhere. Some plants 
usually restricted to high altitudes and open tundra habitats 

224 Rhodora [Vol. 76 

are occasionally found below timberline, e.g. on talus and 
rock piles in Carter Notch (a cool place) and even in some 
open pastures (where they may be rather temporary resi- 
dents). They cannot tolerate shade or root competition of 
lowland plants. A small colony of Mountain Cranberry is 
found along the Half Way House road on Monadnock (pos- 
sibly introduced from the summit by berry pickers). 

However, most steep rocky mountains that, like Monad- 
nock, do not reach timberline and have not been burned 
recently, are densely forested with spruce and at higher 
levels, fir, with a heavy ground cover of moss, ferns and 
herbs. There are probably hundreds of such mountains in 
the Adirondacks and New England. Some of them have 
exposed rocky cliffs, but records showing these to be havens 
for alpine plants in undisturbed condition are few or want- 
ing, Pease (op. cit.) reported finding some arctic-alpines 
on open ledges on secondary mountains in the White Moun- 
tains, *often caused by the destruction of the peaty humus 
by fires," but he did not specify just how often these cases 
were, nor did he give examples of such cases where there 
was no trace or history of fire, He ventured no theory as 
to how these plants reached the lowland habitat. 

If arctic-alpine plants persisted in refugia in lower moun- 
tains after the latter became forest-covered, it should be 
possible to find these plants on unburned summits. They 
have not been found, aside from the cases cited by Pease, 
which were all close to high mountains supporting stable 
colonies of these plants, whence invasion of even small 
burned areas would be relatively easy compared to Mt. 
Monadnock, which is far more remote. 

More examples of bona fide refugia from ice-age times 
need to be found, but even if these can be discovered and 
dated, it may be impossible to demonstrate that no fires 
occurred very long ago, or to prove how long the plants 
have been on the site. Arctic flora of the Pleistocene became 
adapted to the prevailing habitat conditions (Cf. Fernald, 
1925) and it is still largely restricted to areas of open 
ground and a cool, moist climate. 

1974] Flora of Mount Monadnock — Baldwin 225 

Did Arctic-alpine Plants Survive Fire and Spread 
Over the Denuded Areas? 

Assuming that these plants had succeeded in resisting 
the competition of shade-tolerant trees and other plants, 
and had remained in some secluded pocket since post-glacial 
times, did they then survive the very intense fires that 
burned these peaks, more destructive than any fires in 
modern times? Fire destroyed all organic matter, and what 
little mineral soil remained was then at the mercy of wind 
and water. It must be remembered that two great fires on 
Monadnock combined to make a tremendously hot and ex- 
tensive burn. The 1800 fire consumed the humus and killed 
the trees. The 1815 hurricane completed toppling them, so 
that a formidible quantity of dry fuel was present. (It was 
after a long hot, dry summer.) No fires in recent times 
have approached it in heat and ferocity. Emerson (1896) 
remarked that the heat was so great that it “blasted the 
rock" into the ravines and caused slides. 

How did Arctic-alpine Plants Come to Occupy Areas 
Denuded by Fire? 

Such plants must have arisen from disseminules brought 
from nearby areas (refugia on the same mountain or in the 
lowland) or from more distant sources, probably stretches 
of alpine tundra on Mt. Moosilaukee or on the Presidential 
Range. Since the lowland and lower slopes around Monad- 
nock were largely cleared and cultivated or pastured, it is 
possible that some of these plants had escaped from rocky 
crevices on the mountain before the fire and had become 
established on the lower open land. Here they might main- 
tain themselves until forest succession once more eliminated 
them. At the time of the fire, this land was still open. Even 
if restocking of the burned mountain had proceeded from 
this source, it is hard to see how the spread of heavy-seeded 
species could have occurred uphill without the aid of birds 
or animals. The latter carry seeds in burs that cling to 
animal fur, but animals have not been found to be effective 
carriers of berries and similar hard seeds. 

226 Rhodora [Vol. 76 

Were Birds Partly Responsible for the Spread of Plants 
From Alpine Tundra to Monadnock? 

Birds are well-known to be active and efficient carriers 
of seeds. Ridley (1930) noted that birds are attracted by 
seed color. Red seems to be the most attractive. Vaccinium 
is a “bird-dispersed genus", he wrote. Forty-three species 
of birds have been recorded to disperse plants of this genus, 
and it seems likely that the red fruit of Mountain Cranberry 
would be very much sought by birds. Taylor (1954) sug- 
gested that birds are probably responsible for carrying 
seeds to Macquaric Island, a small sub-antarctic island far 
from any large land area that was severely glaciated during 
the Pleistocene. He believed it unlikely that any of the 35 
plant species could have survived the ice. Most of these 
have relatively large seeds, but all have propagules suited 
to bird transport. 

Birds are important agents in the natural dissemination 
of seeds of tree species having fleshy fruits, and presum- 
ably shrubs as well. Red Cedar, Black Cherry and Black 
Gum are examples. Seeds of Juniperus that have been 
eaten by birds are said to germinate more promptly. (Anon. 
1948) Indeed such hard-seeded species that display seed 
coat dormancy may require the acid treatment received 
during passage through a bird’s intestinal tract in order to 
germinate. Phillips (1910) made an especially thorough 
and wide-ranging study of bird dissemination of Juniperus 
seed, and concluded that birds are responsible for most of 
the spread of junipers, especially upwards on slopes and 
up stream courses. 

It is possible that the most likely candidate for being a 
vector of seeds from alpine fell fields is the Slate-Colored 
Junco. Not only did Thoreau remark about its being the 
most common bird on Monadnock, but it is also abundant 
there today. Forbush (1929) states that it is probably the 
only bird that nests above tree limit in the White Moun- 
tains. (I have found a nest with eggs in late June on Boott 
Spur.) Forbush (op. cit. p. 90) reported that it eats many 

1974] Flora of Mount Monadnock — Baldwin 227 

dried berries and seeds. Objections to the theory of bird 
transport include the questions concerning birds eating be- 
fore migration, seeds passing undamaged, and limits set by 
the time to pass through the digestive tract. Egler (1972) 
questions the explanation “by having far-flying constipated 
birds eject over distant lands.” 

While there is thus evidence that some birds do carry 
seeds, and birds are present on Mt. Monadnock, and that 
seeds survive the treatment, or may be even benefited by 
it, it is still not proven that heavy-seeded arctic-alpine 
plants did actually become established on Monadnock by 
this means. 

Since this was written, Steele and Hodgdon (1973) have 
examined these theories for the arrival on Mt. Cardigan of 
two alpine species of Carex. They suggest a third possibil- 
ity, that man may have consciously or unconsciously intro- 
duced these plants to the bare rocky area. This appears 
least likely in the case of the plants found on Monadnock, 
because of the distance. It is also improbable that these 
plants should appear on Mt. Chocorua and other sites. It 
would be a very diligent Johnny Appleseed indeed. More 
important, a prolonged period must be assumed for estab- 
lishment. Success probably did not come with the first seed. 

I am very grateful to Dr. Albion R. Hodgdon for checking 
the identity of specimens collected, and for helpful sugges- 
tions on the manuscript, also to Mr. David E. Boufford for 
searching herbaria and listing Monadnock records. 


ANON. 1948. Woody-Plant Seed Manual. Misc. Pub. 654, U.S. Dept. 
Agri. 416 p. 

BRAUN, E. Lucy. 1950. Deciduous Forests of Eastern North Amer- 
ica. Blakiston, 596 p. 

CHAMBERLAIN, ALLEN. 1936. Annals of the Grand Monadnock. 
SPNHF, Concord, N.H. 195 p. 

CHILD, HAMILTON. 1885. Gazetteer of Cheshire County, N.H. 1736- 
1885. Pub. by the Daily Journal, Syracuse, N. Y. 560 p. 

CUTTER, DANIEL B., M.D. 1881. History of the Town of Jaffrey, 
N.H. Concord, N.H. 648 p. 

228 Rhodora [Vol. 76 

DANA, JAMES FREEMAN. 1816. Some Account of the Grand Monad- 
nock. New England Jour. of Medicine and Surgery 5: 252-257. 
Boston, Mass. 

DEANE, WALTER. 1890. Flora of the Summit of Mt. Monadnock, 
N.H. in July. Bull. Torr. Bot. Club 17: 316-318. 

EGLER, FRANK E. 1972. (in a review) Ecology 53(5): 988. 

ELLIS, GEORGE B. 1880. Memoir of Jacob Bigelow. Proc. Mass. Hist. 
Soc. 17: 383-467. 

EMERSON, EDWARD W. 1896. The Grand Monadnock. New England 
Magazine 15: 33-51. 

FERNALD, M. L. 1907. The Soil Preferences of Certain Alpine and 
Subalpine Plants. Rhodora 9: 149-193. 

1925. Persistence of Plants in Unglaciated Areas 

of Boreal America. Memoirs Am. Acad Arts & Sciences 15: 235- 


. 1950. Gray's Manual of Botany. Eighth Edition. 
Am. Book Co., N.Y. 1632 p. 

ForBUSH, Epwarp Hower. 1929. Birds of Massachusetts and other 
New England States. Part III. Mass. Dept. of Agri. Norwood, 
Mass. 466 p. 

LEONARD, LEVI W. 1855. History of Dublin, New Hampshire. Bos- 
ton, Mass. 433 p. 

PEASE, A. S. 1964. A Flora of Northern New Hampshire. New 
England Botanical Club. Cambridge, Mass. 278 p. 

PHILLIPS, FRANK J. 1910. The Dissemination of Junipers by Birds. 
Forestry Quarterly 8: 60-73. 

RIDLEY, HENRY N. 1930. The Dispersal of Plants Throughout the 
World. L. Reeve & Co., Ashford, Kent, England. 744 p. 

Spurr, S. H. 1964. Forest Ecology. Ronald Press, N.Y. 352 p. 

STEELE, F. L. and A. R. Hopepon. 1973. Two Interesting Plants on 
Mt. Cardigan. Rhodora 75: 155-157. 

STONE, WILLIAM H. 1885. The Flora of Mt. Monadnock. Appalachia 
4: 145-150. 

TAYLOR, R. W. 1954. An Example of Long Distance Dispersal. 
Ecology 35: 569-572. 

THOREAU, HENRY D. (1906) 1962. The Journals of Henry D. Tho- 
reau 1852-1860. Edited by Bradford Torrey and Francis H. Allen. 
14 Vol. bound as two. Dover Pub. N.Y. 




Section PERITYLE, with 27 species and three varieties, is 
the largest of three sections of the genus Perityle (Powell, 
1968a). The distributional center of sect. PERITYLE is in 
northwestern Mexico and the Baja Californian peninsula, 
but the total natural range is extended by several taxa that 
occur in the southwestern United States, Guadalupe Island, 
the Revillagigedo Islands, and west-central Mexico (Jal- 
isco). One weedy taxon, P. emoryi, has bicontinental dis- 
tribution in North America and in Chile and Peru in South 
America. Separate taxonomic works have been prepared 
for the other two sections of the genus; Pappothrix (Powell, 
1969) and Laphamia (Powell, 1973). 

The species of sect. PERITYLE comprise a taxon that was 
recognized as the genus Perityle by Everly (1947) who 
compiled a taxonomic treatment of the group. In connection 
with her morphologically oriented investigations of Perityle, 
Everly also studied the closely related taxa Pappothrix and 
Laphamia, which she regarded as genera. These taxa are 
distinguished primarily by somewhat variable pappus and 
achene characteristies, and several workers have questioned 
their status as genera. Shinners (1959) merged Pappothrix 
and Laphamia with Perityle, noting their close morpho- 
logical relationship and contending that pappus differences 
were not sufficient to warrant separate generic status. In 
recognizing Pappothrix and Laphamia as sections of Peri- 
tyle, I have followed Shinners’ congeneric disposition of 
the taxa (Powell, 1968a). My systematic judgements were 
based upon the evaluation of new information from cyto- 
logical, chromatographic, and hybridization techniques, as 
well as a re-evaluation of morphological] features. The re- 

'Supported by National Science Foundation Grants GB-7740 and 


230 Rhodora [Vol. 76 

sults have suggested that Pappothrix, Laphamia, and Peri- 
tyle are closely related, essentially natural phylads, and 
that they are best treated as sections of a single genus. 
Niles (1970), however, has maintained Perityle and Lapha- 
mia as distinct genera, while combining Pappothrix with 
Laphamia. Niles did not conduct a comprehensive study of 
the sect. PERITYLE species. A more complete review of the 
taxonomic history concerning Perityle is available elsewhere 
(Shinners, 1959; Powell, 1968a; Niles, 1970). 

Perityle is closely allied to Pericome, Amauria, and 
Eutetras of the subtribe Peritylinae (Helenieae). Rydberg 
(1914) assigned Perityle and Pericome to the Peritylinae 
and Amauria and Ewutetras to the subtribe Amaurinae. 
Turner (1966) and Powell (1972a) have suggested that 
Amaurinae (in part) should be combined with Peritylinae. 
Presently the Peritylinae is being re-evaluated with refer- 
ence to Rydberg's concepts, and the proper tribal position 
of the subtribe is also being investigated (Powell & Turner, 


A discussion of the chromosome numbers of Perityle and 
related genera is presented elsewhere (Powell, 1968b). 
Additional counts for the species of sect. PERITYLE are 
reported in Powell and Sikes (1970) and Powell (1972b). 
Chromosome numbers have been obtained for 20 of the 27 
species of sect. PERITYLE. The counts for specific taxa are 
included with a list of the species under Phylogenetic 
Considerations (Table 1), and they are also included with 
species discussions in the taxonomic portion of this paper. 

In Powell (1968b), a few changes must be noted with 
regard to the counts for sect. PERITYLE. When the 1968 
paper was compiled, the taxonomic studies of sect. PERITYLE 
were in early stages. The reports of n — 16 for P. cali- 
fornica should be attributed to P. cuneata var. marginata. 
Perityle incompta is now considered as synonymous with 
P. crassifolia var. robusta, and the counts listed for the 
former (n = 19, 18) should be attributed to the latter 

1974] Perityle — Powell 231 

taxon. The diploid P. microglossa (Sikes and Babcock 185 
a,b,c,f,) is P. microglossa var. saxosa, and the polyploid 
P. microglossa (Sikes and Babcock 177 192 a-d, 152, 157 a, 
206 a,b,) is var. microglossa. Sikes and Babcock 188 a,b,d,f, 
and 190 a,b, listed as P. palmeri, are P. cordifolia (n = YT). 
Perityle cf. spilanthoides is P. turneri (n — 1T) and P. 
vaseyi (Sikes 116) is P. parryi (n — YT). 

Most species of sect. PERITYLE are diploid, with counts of 
n — 19, 18, 17, 16, 13, 12, and 11 having been reported for 
the various taxa (Powell, 1968b; 1972b). Only P. micro- 
glossa var. microglossa (n = 34, 51), P. emoryi (n = 32-36, 
50-57), and P. incana (n = 50-57) are known to be poly- 
ploid. The first two taxa are relatively well established as 
exclusively polyploid, while only one count for the Guada- 
lupe Island endemic P. incana (Powell, 1972b) has been 
recorded. Turner and Flyr (1966) reported n = ca. 46 for 
P. crassifolia var. robusta (Cowan 2265), but I have since 
identified this collection as P. emoryi. A base number for 
P. emoryi has not been determined in spite of numerous 
attempts to obtain exact counts (Powell, 1968b). 

The ancestral base number for sect. PERITYLE has not 
been established. Subsequent to an earlier discussion of 
the subject (Powell, 1968b), all attempts to resolve the 
question have been unsuccessful, including those which in- 
volved the analysis of experimenta] hybrids (Powell, un- 
published). Most chromosomal] and distributional evidence 
suggest a base of x — 17 for sect. PERITYLE, and the genus 
as a whole (see Phylogenetic Considerations). Fourteen 
species of sect PERITYLE are n = 17, and both sect. PAP- 
POTHRIX (5 spp.) and sect. LAPHAMIA (21 spp.) are x = 17 
(Powell, 1969; 1973). The taxa of sect. PERITYLE with 
chromosome numbers below n = 17, P. californica (n = 13, 
12, 11) and P. cuneata (n = 16, 12), are clearly aneuploid 
derivatives, probably of the » — 17 line. However, the 
origin of taxa with n = 19 (P. crassifolia and allies) is of 
prime concern in evaluating the phylogeny of sect. PERI- 
TYLE. Distributionally, the n = 19 “alliance” ( (southern 
Baja California peninsula and neighboring islands) is pe- 

232 Rhodora [Vol. 76 

ripheral to the group where n — 17, which is centered on 
the mainland of Mexico (see Phylogenetic Considerations). 
The » — 19 group may be either ancestral to or derived 
from the » — 17 group (Powell, 1968b), but I now believe 
that the species with » — 19 and the species with » — 17 
both evolved as separate lineages from a common “main- 
land" ancestor that had a chromosome number of n = 19 
(or 18). Consideration of the latter hypothesis in a broader 
sense is supported by the base numbers of the related 
genera Pericome (x = 18), Eutetras (x = 18), and Amau- 
ria (x = 18). 


Artificial hybridizations have been conducted with 16 
species of sect. PERITYLE. Many of the crosses were of 
intergeneric and intersectiona] nature, although several 
interspecific combinations have been obtained. The results 
of most of these crosses are discussed in another paper 
(Powell, 1972c) where emphasis was placed upon evalua- 
tion of the intergeneric and intersectional relationships of 
Perityle. A hybridization program aimed at the evaluation 
of interspecific relationships within sect. Perityle is still 
in progress (Powell, unpublished). Where appropriate, 
some of the preliminary information derived from artificial 
crosses will be brought into species discussions in the taxo- 
nomic portion of this paper. 

Natural hybridization is rare in sect. PERITYLE, and only 
one such occurrence has been documented (Powell, 1970). 
Most of the species are geographically isolated, but a few 
taxa do have overlapping ranges and may occasionally hy- 


Nineteen species of sect. PERITYLE were analyzed by 2- 
dimensional paper chromatography of leaf extracts. In 
general, the techniques outlined by Alston and Turner 
(1963) were utilized. The procedures for sampling popu- 

1974] Perityle — Powell 233 

lations and analyzing chromatograms have followed those 
discussed elsewhere (Powell, 1973). The major components 
of chromatographic patterns were characterized as flavo- 
noids (Powell and Tsang, 1966; Powell and Averett, un- 
published), although the specific identity of only one com- 
pound is known (Southwick et al., 1972). 

The use of chromatographic techniques has been a valu- 
able asset to previous taxonomic studies in the Peritylinae 
(Powell and Tsang, 1966; Powell, 1968a; 1969; 1973). 
Intergeneric and infrageneric relationships have been clari- 
fied through the comparative analysis of pattern data. In 
previous work I treated the crude chemical information 
from a chromatographic pattern as a single taxonomic 
character, and pattern data have not been given more 
weight than any other chraacter. 

The results of chromatographic studies in sect. PERITYLE 
have revealed two strikingly different chemical patterns 
based upon the presence or absence of yellow compounds 
as seen under ultraviolet light. The white-flowered species 
centered in the Sierra Madre Occidental (e.g., P. turneri), 
and the yellow-flowered species of northwestern Mexico and 
Baja California (e.g., P. cordifolia and P. californica) have 
simple patterns consisting of a few dark purple spots. The 
Baja Californian white-flowered species (e.g. P. crassi- 
folia), and the Texas yellow-flowered species (e.g., P. par- 
ryi) have relatively complex patterns that include promi- 
nent yellow spots. Yellow compounds are considered im- 
portant because they comprise the basic chromatographic 
profile of sect. PAPPOTHRIX, sect. LAPHAMIA, and the genera 
related to Perityle. The actual chromatographic data are 
not presented here in the form of figures or tables. Instead, 
where appropriate, the data are included in succeeding dis- 
cussions concerning relationships of species and species 


The species of sect. PERITYLE occur in rock and in soil 
habitats, unlike the species of sect. PAPPOTHRIX and LAPHA- 

234 Rhodora [Vol. 76 

MIA, which are exclusively rock-dwelling (Powell, 1969; 
1973). Certain species of sect. PERITYLE are obligate, rock- 
dwelling perennials, while several perennial and annual 
taxa are soil-obligates. In addition, a few taxa display 
facultative existence in rock and soil habitats. 

The rock-dwelling members of sect. PERITYLE occur under 
a wide range of climatic conditions and at various eleva- 
tions. About half of the petrophilic taxa are occupants of 
protected canyons and exposures in relatively xeric desert 
mountains at altitudes of 1000-4000 feet or less, and the 
others thrive under relatively mesic conditions at altitudes 
of 4000-8000 feet. Perityle turneri also occurs under mesic 
conditions, at 7000-9500 feet in and near Durango, Mexico, 
but may be rooted primarily in matted soils. The other soil- 
dwelling taxa occur almost exclusively under desert condi- 

Perityle is essentially a genus of self-incompatible spe- 
cies (Powell, 1972c). Only two of the taxa, P. emoryi and 
P. microglossa var. microglossa, are known to be self-com- 
patible, and these are members of sect. PERITYLE. Ten spe- 
cies have not been tested for the compatibility factor, two 
of sect. LAPHAMIA and eight of sect. PERITYLE, but these 
taxa are also probably self-incompatible. Both of the self- 
compatible taxa are widespread polyploids (Powell, 1968b; 
1972c) that are “weedy” and annual in habit. Perityle 
emoryi is the most widespread species of the genus, having 
bi-continental distribution in North and South America, 
and P. microglossa var. microglossa probably ranks second 
in distributional range. Several other species of the genus 
are polyploid, and a few others are annual, thus suggesting 
that the capacity for reproductive self-compatibility has 
allowed for the “weedy” nature of two taxa. 


Speculation concerning phylogeny of the subtribe Perity- 
linae and the genus Perityle has been discussed generally 
elsewhere (Powell, 1973), and will not be repezted in detail 
here. Essentially, it was proposed that Peritylinae origi- 

1974] Perityle — Powell 235 

nated in the southern Sierra Madre Occidental of Mexico. 
Perityle is by far the most successful genus of the subtribe, 
in terms of number of species, with only 2 or 3 species being 
recognized for each of the other genera (Powell, 1972a; 
1972d; and Turner, 1966). Section PERITYLE is assumed to 
be ancestral to the other two sections of the genus. Based 
upon distributional, morphological, chromosomal, and other 
considerations, it is believed that sects. PAPPOTHRIX and 
LAPHAMIA underwent rapid evolution in late Pliocene or 
Pleistocene emanating from an ancestral sect. PERITYLE 
complex (Powell, 1975). 

Modes of speciation in the subtribe Peritylinae, and Peri- 
tyle in particular, have been discussed elsewhere (Powell, 
1969; 1972c; 1973). In sects. PAPPOTHRIX and LAPHAMIA, 
speciation is believed to have occurred mostly, if not en- 
tirely, as a result of geographic isolation. In sect. PERITYLE, 
speciation probably has occurred as a result of geographic 
isolation and other, more nearly reproductive phenomena 
(Powell, 1972c). This subject wil be diseussed further 

In Table 1 the list of sect. PERITYLE species is presented 
as an aid to discussing phylogenetic considerations. The 
list has been organized to depict the hypothetical evolution- 
ary groupings within the section. The arrangements are 
based upon morphological, chromosomal, experimental, 
chromatographic, and eco-geographical data. Additional 
discussions of species relationships, including the evidence 
upon which such judgements are based, are included in the 
taxonomic portion of this paper. 

The total data suggest that sect. PERITYLE comprises at 
least two distinct evolutionary series. One series, including 
the first 19 taxa, is marked by white ligules and yellow (or 
cream-white) disc corollas, with a few exceptions, as indi- 
cated in Table 1. Perityle incana and the P. jaliscana group 
are relatively anomalous in the “white-flowered series" and 
they may comprise separate and equal phyletic entities. A 
second major series can be delimited by its exclusively 
yellow flowers. 


[ Vol. 



‘SNOGUOLLI Ə OF JYSNOY} ƏV jnq *exej eArjedsei ay}, J04 pojiodet useq eAvY seseyjueded ur s.9quin N.. 



(61) LI 149w]pnd'q “GZ 
LI 04000) 'd "yc 
pe1unoo 4ou uphüol]'d ec 
LI 0$$0]D02d9]'d ZZ 
LT )050f1p402'd "Ic 
LI (pioosrp) 08s0]bD'd  '0c 
(6I) LI Wiosna'q ‘61 
LI Wia4Dnd, ^q “RT 


pojunoo jou (proosrp) omuoppaí'qd "LI 
pejunoo 30u DJUOPUOYNA ‘d  '9[ 
p91unoo jou 19804 `d GI 
LI (proosrp) pounosqp('q ‘I 

ISI 93IQA-urga4?) pue Se INYM 

LT « (91) 

po1unoo jou 

Ic ‘FS 

peyunod jou 
peyunod jou 
LT “89 ‘LT 

L€-0€ ‘9E-ZE 


D1423$19WJ0 “dq “ST 
nyofidouoLod *q ‘ZT 
0401% 'q “TT 
0D90]1.409W1] `A 
149W402 *d 
(proosrp) mwnDow'q 
(SABI Mo[[oÁ) 02470 “gq 
D10f1$8049 `A “QT 


Aled Has SOs 

‘sdnois seroeds-peje[or peumnsaid puge 1009 IƏMOY 0} SurpJooo*€ PASI ext, 

sIequinu eurosouroIq9 orjeure3 jo Áueurums put seroeds AÁ uoreg 


1974] Perityle — Powell 237 

Several related-species groups are further delimited 
within both the ‘“white-flowered series" and “yellow- 
flowered series” (Table 1). The groupings themselves com- 
prise clearly related taxa, but the phyletic unity of the 
related-species groups within monophyletic white- or yellow- 
flowered series is indefinite. For example, origin of the 
Baja Californian P. crassifolia group has not been con- 
nected resolutely with the Sierra Madre Occidental P. 
turneri group, even though both exhibit white ligules. And 
the anamalous nature of P. incana and the P. jaliscana 
group is indicated above. Also, there is no certainty that 
the P. parryi group of Texas and the aneuploid P. cali- 
fornica group of Baja California have monophyletic ties 
within a yellow-flowered series. 

White-Flowered Groups (Table 1). Three subgroups of 
the white-flowered series are recognized. These are desig- 
nated as the Perityle crassifolia, P. turneri, and P. jaliscana 
related-species groups. Distributional information, habitat 
preferences, and life form habits are given in the following 
diseussions because of their supposed importance in under- 
standing speciation in the genus. 

The Perityle crassifolia group, also including P. socor- 
rensis, P. emoryi, and P. aurea, is distributed in southern 
Baja California and nearby islands, except for P. emoryi 
which is a widespread weed. Also, P. socorrensis is found 
only in the Revillagigedo Islands, which are over 200 miles 
south of Baja California. 

Perityle emoryi (n = 32-36, 50-57) is believed to be a 
polyploid derivative of P. crassifolia var. robusta. Morpho- 
logical and chromatographic data strongly suggest that 
P. aurea belongs with the P. crassifolia (x — 19) group 
even though this taxon has yellow-ligules and a chromo- 
some number of n = 17. 

Both Perityle emoryi and P. aurea are annual in habit, 
while the other taxa of the P. crassifolia group are essen- 
tially soil-dwelling perennials. All of the taxa, however, 
with the exception of P. crassifolia var. crassifolia which 
seems restricted to sand, might also occur in soil-filled 

238 Rhodora [Vol. 76 

crevices of rocks especially at sea side. The Guadalupe 
Island endemic P. incana, is possibly allied with the P. 
crassifolia group, but evidence as to its true relationship 
has not been conclusive. 

The base chromosome number of the Perityle crassifolia 
group is presumed to be x — 19, a number that might be 
ancestral in sect. Perityle. On morphological and distribu- 
tional grounds, however, it is practically impossible to en- 
vision the other related species groups as having been de- 
rived from the P. crassifolia line. I believe that the P. 
crassifolia line was an early offshoot from ancestral Peri- 

The largest related-species group of the white-flowered 
series is distributionally centered in the Sierra Madre 
Occidental] of mainland Mexico. Two assemblages of four 
species each are further distinguishable among this Perityle 
turneri group, primarily on the basis of floral characters. 
Perityle turneri, P. microcephala, and P. lineariloba are 
clearly allied taxa which occur at relatively high altitudes 
from Durango to central Chihuahua. The fourth member 
of this assemblage is P. microglossa, which ranges from 
northern Sonora along the west coast of Mexico south to 
Colima, into the foothills of the Sierra Madre Occidental, 
and east to San Luis Potosi and Texas. Habit and distribu- 
tional considerations (see Taxonomy) strongly suggest that 
P. microglossa is derived from P. microcephala. I believe 
that P. turneri is primitive in this assemblage, on the basis 
of features discussed elsewhere (Powell, 1972e). 

Perityle coronopifolia, P. ciliata, P. hofmeisteria, and P. 
canescens are also clearly allied members of the P. turneri 
group. Perityle coronopifolia extends from northern Chi- 
huahua into the mountains of southern Arizona and south- 
ern New Mexico. Perityle ciliata is restricted to central 
Arizona, while P. hofmeisteria and P. canescens apparently 
are endemics respectively in Durango and in Sinaloa, 

Most members of the Perityle turneri group are hardy, 
suffrutescent perennials that live in crevices of rock bluffs. 

1974] Perityle — Powell 239 

Perityle turneri is a rhizomatous perennial, apparently 
occurring typically in thin soil covering rocky outcrops, but 
according to label data, the taxon might also be rock-dwell- 
ing. Perityle microglossa is a soil-dwelling annual, also 
found occasionally among rocks, and one collection of P. 
hofmeisteria (Flyr 332) is said by the collector to be “seem- 
ingly an annual.” Judging from known distributional data, 
these species are strictly allopatric, suggesting that they 
have undergone the type of geographic speciation that is 
considered typical for sect. PAPPOTHRIX and sect. LAPHAMIA 
(Powell, 1969; 1973). 

The white-rayed (or discoid) Perityle jaliscana group 
with cream-white discs comprises very closely related rock- 
dwelling perennial taxa that are restricted to Jalisco, Mex- 
ico. These taxa represent a third distinct assemblage within 
the white-flowered species of sect. PERITYLE. 

The three white-flowered subgroups are clearly delimited 
morphologically, with perhaps the most useful distinguish- 
ing features being those of the habits, achenes, and pappus 
(see Taxonomy). The yellow-flowered species are quite 
different superficially, but the generic unity of both white- 
and yellow-flowered assemblages is evident. 

Yellow-Flowered Groups (Table 1). Geographically, the 
yellow-flowered species are divided into two major assem- 
blages separated by the Sierra Madre Occidental. One as- 
semblage, designated as the Perityle parryi group, has a 
limited distribution in the Big Bend area of western Texas 
and adjacent Mexico, and the other geographic assemblage, 
including the P. cordifolia and P. californica groups, is 
more widely distributed in Sonora, Sinaloa, and Baja 
California, Mexico. 

The three species of the Perityle parryi group are closely 
related perennials, being distinguished on the basis of a 
few morphological characters. One of the taxa, P. aglossa, 
apparently is exclusively rock-dwelling in habit, while P. 
vaseyi seemingly occurs only in desert soils. Perityle parryt, 
however, is facultative in its existence in rocks and soils. 

There are interesting biological parallels between the 

240 Rhodora [Vol. 76 

Perityle parryi group and the P. cordifolia group of west- 
ern Mexico, and despite their geographic separation, most 
evidence suggests that both groups had common origin. 
Both alliances comprise closely related taxa, their vegeta- 
tive and floral morphologies are strikingly similar, and 
species of both groups have chromosome numbers of n = 17 
with no polyploidy having been discovered in any of the 
taxa. Additionally, and most remarkably, the evolutionary 
pattern of the disjunct groups seems to be that soil forms 
were established when woody perennials “came down out 
of the rocks and adapted to life in the soil.” Perityle cordi- 
folia, P. leptoglossa, and P. lloydii seem to be obligately (at 
least typically) rock-dwelling, while P. palmeri and P. 
lobata apparently are facultative in their tolerance for 
existence in rocks or in soils. 

The presumed “close” phylogenetic relationship between 
the Perityle parryi and P. cordifolia groups is not supported 
by their chemical profiles. Members of the P. parryi group 
produce abundant yellow components, as do P. crassifolia 
and allies, while chromatographic samples of the P. cordi- 
folia group have revealed only simple patterns of purplish 
spots. The chemical indication of affinity between the P. 
parryt and P. crassifolia groups is complemented by a 
tenuous morphological feature they have in common. Peri- 
tyle vaseyi (rarely P. parryi, also) of the former alliance 
and pappose members of the P. crassifolia group are the 
only taxa of the genus which have awns with retrorsely- 
barbed tips. In view of the morphological dissimilarity 
and geographic distance between these groups, however, I 
believe that the crude chemical data and pappus structure 
merely reflect an ancestral connection between the P. crassi- 
folia and yellow-flowered alliances. If this is true, then the 
P. cordifolia group must have lost its ability to produce the 
prominent yellow flavonoids. 

Both species of the Perityle californica group are soil- 
dwelling annuals. The taxa are of special systematic inter- 
est in sect. PERITYLE because of their reduced chromosome 
numbers. All other yellow-flowered taxa have m — 17. 

1974] Perityle — Powell 241 

Perityle californica (n = 13,12,11) is distributed in Sonora 
and Sinaloa of mainland Mexico and along most of the Baja 
California peninsula to just south of La Paz. Perityle 
cuneata (n = 16,12) is restricted to the southern tip of 
Baja California with northern limits to about the latitude 
of La Paz. 

Judging from overall morphological similarity and dis- 
tributional considerations, it is assumed that Perityle cali- 
fornica and P. cuneata are aneuploid derivatives of the P. 
cordifolia line. With the exception of the P. parryi group, 
which seemingly would be ruled out on distributional 
grounds, only the P. cordifolia group bears any close re- 
semblance to the aneuploids. Perityle californica occurs 
sympatrically with P. cordifolia and P. palmeri in Sonora 
and Sinaloa, apparently without hybridization. Since most 
species of Perityle are allopatric, but artificially interfertile 
(Powell, 1972c), it is reasonable to conclude that P. cali- 
forniea achieved reproductive isolation and speciation 
through reduction in chromosome number. 

Summary of Phylogenetic Considerations. Species of the 
genus Perityle are distributed primarily in the arid to semi- 
mesic mountains of desert North America, but also in the 
higher mountains, for example in the Sierra Madre Occi- 
dental. The majority of the species are obligate rock- 
dwellers. All five species of sect. PAPPOTHRIX and all 21 
species of sect. LAPHAMIA occur as perennials in crevices 
of rock bluffs, apparently never growing in soil at the base 
of bluffs. That these species can survive only in such re- 
strictive habitats surely imposes upon them evolutionary 
limitations. The basic evolutionary trends of sect. PAPPO- 
THRIX and LAPHAMIA have followed a pattern of speciation 
by geographic isolation; thus evolved the groups of closely 
related, mostly endenmic species, characteristic of the above 

Greater evolutionary diversity is exhibited among the 
species of sect. PERITYLE. Indeed, some species of this sec- 
tion are obligate, rock-dwelling perennials that presumably 
underwent the type of geographic speciation mentioned 

242 Rhodora [Vol. 76 

above. But several taxa have adapted to a facultative ex- 
istence in rocks and in soils, and a few species are exclu- 
sively soil-dwelling with perennial or annual habits. For 
certain taxa, the adaptation to life in the soil apparently 
has been facilitated by two evolutionary mechanisms, poly- 
ploidy and aneuploidy. Emanating from the white-flowered 
groups, the widespread weeds Perityle microglossa var. 
microglossa (n = 34,51) and P. emoryi (n = 32-36, 50-57) 
exemplify the adaptive success of polyploidy. Perityle cali- 
fornica (n = 11, 12, 19) and P. cuneata (n = 12, 16) are 
successful aneuploid species of yellow-flowered lineage. 
Thus, along with the annual habit displayed by the above 
species, the phenomena of polyploidy and aneuploidy have 
independently advanced the adaptive capacity of species 
belonging to distinct evolutionary units of sect. PERITYLE. 


I am grateful to the curators of herbaria from which 
specimens were borrowed. I am also grateful to B. L. 
Turner who suggested the study of Perityle and provided 
encouragement and advice throughout the study. I extend 
thanks to John Averett for his assistance in gathering 
literature; to Reid Moran for his help in providing vouch- 
ers, buds, and ecological information for P. incana and 
other species; to Rogers McVaugh for providing helpful 
observations and material of P. microglossa and P. fedde- 
mae; and to Annetta Carter for her assistance in providing 
specimens and ecological information for several species. 
Many other people have contributed to my studies of Peri- 
tyle and related genera; they know of my appreciation. 
One person who did not know fully of my deep gratitude 
was the late Lloyd Shinners. His special character will 
linger to inspire many taxonomists. 


PERITYLE Benth. Bot. Sulph. 23. 1844. 
Perityle section Perityle, see Powell, Sida 3: 277. 1968. 

1974] Perityle — Powell 243 

Galinsogeopsis Schultz-Bip. in Seem. Bot. Herald. 306. 1856. 
Nesothamnus Rydb. N. Amer. Fl. 34: 12. 1914. 
Leptopharynz Rydb. N. Amer. Fl. 34: 21. 1914. (in part). 
Closia Remy in Gay, Fl. Chile 4: 120. 1849. (in part). 

Plants shrubby, suffruticose, or herbaceous perennials, 
or delicate to robust annuals, (2)10-80 cm high, growing 
in rock crevices or various soils, variously pubescent, to- 
mentulose to nearly glabrous, frequently glandular-pu- 
bescent; leaves opposite or alternate, variable in size and 
shape, 0.7-14 cm long including the petiole, 0.2-7.0 cm wide, 
ovate, cordate, deltoid, subhastate, reniform, or suborbicu- 
lar in outline, the margins subentire, serrate to doubly so, 
or shallow to deeply lobed or divided, frequently 3-5 lobed, 
pedate, subcruciform, or pinnatifed with narrow segments, 
nearly glabrous, puberulent, to arachnoid-villous, often 
glandular-pubescent, thick and semisucculent to thin in 
texture, viscid, bitter tasting, usually aromatic; capitu- 
lescence of solitary heads, or heads loosely to tightly aggre- 
gated in clusters of 3 or more; peduncles short or long; 
involucres subcylindric to hemispheric; heads 3-12 mm high, 
4-15 mm wide; involucres of 2(3) subequal series, bracts 
ovate or obovate to narrowly oblanceolate, obtuse to at- 
tenuate at the apex, flattened, ribbed, or keeled, usually 
spreading or reflexing at maturity; heads radiate or discoid 
(in 4 spp.), ligules yellow or white, showy or rudimentary ; 
disc flowers numerous, corollas 4-lobed, yellow or cream- 
white (in 3-4 spp.), throats tubular to broadly funnelform 
or narrowly campanulate; style branches, flattened, linear, 
usually tapering to a fine, minutely pubescent tip; achenes 
black when mature, 1.3-5.0 mm long, distinctly flattened 
radially, outer achenes often 3-angled, slightly curved and 
clasped by bracts, linear, oblong, oblong-elliptic, or oblance- 
olate, the margins densely ciliate, rarely merely puberulent, 
prominently calloused or the margins thin and not cal- 
loused, the faces glabrous and shiny or short-pubescent; 
pappus of a conspicuous, rarely inconspicuous, crown of 
laciniate, hyaline, squamellae, and 0-2 (rarely 3-4) delicate 
or rather stout bristles, 0.5-7.0 mm long, the bristles naked, 

244 Rhodora [Vol. 76 

barbellate, or subplumose throughout or only at the tips, 
the barbs antrorse, lateral, or retrorse; base chromosome 
number, x = 17 or 19. 

Type: P. californica Benth. 

Key to the Species 

1. Heads discoid. 2 

a somos otov t 9 9 n nm m m m rs n, n n n n n. n s s n. d 

2. Leaves canescent; Guadalupe Lsland, Mexico. .... 
TEM 5. P. incana. 

2. Leaves not canescent; Texas or Jalisco, Mexico . 3. 
3. Pappus bristle 1, 3.7-5.5 mm long; disc corollas 

yellow; Texas. ................ 20. P. aglossa. 

2 Pappus bristles 2(3), 0.5-1 mm long; dise corol- 

las cream-white or pale yellow; Jalisco. ...... 4. 

4. Leaves dissected. ........ 17. P. feddemae. 

4. Leaves deltoid to subhalberd. 14. P. jaliscana. 

1. Heads radiate. .... ""arzsKAazPe»saerszrresruss D 
5. Ray and disc corollas yellow. ................. 6. 

6. Plants distinctly annual; pappus bristles 1 or 2 

(or rarely absent). ....................... T. 

7. Achene margins thin, not calloused; single 
pappus bristle delicate (rarely absent), and 
retrorsely barbellate only at the tip. ........ 
MM 4. P. aurea. 

7. Achene margins usually prominently cal- 
loused; single pappus bristle rather stout 
and antrorsely subplumose, or 2 delicate 

bristles. .... .0.0............ 8. 
8. Pappus bristle 1, subplumose, 1.5-3.5 mm 
long. ...... 2 ee. 26. P. californica. 
8. Pappus bristles 2 (rarely absent), delicate, 
0.5-2.0 mm long. .................... 9. 

9. Achenes (2.5) 3-3.8 mm long, obcor- 
date-cuneate, with broad, callous mar- 
gins. ... 27a. P. cuneata var. cuneata. 

9. Achenes 1.5-2 (2.5) mm long, obovate 
to subcuneate with prominent callous 


Perityle — Powell 245 

TOTS suc Se a 
27b. P. cuneata var. marginata. 

6. Plants perennial, with woody bases, or herba- 

ceous with fleshy taproots or rather thin branch 

Toota 2 ee ec ele oe 10. 
10. Plants of West Texas and adjacent Mexico. 
ACTI NS DEUS or weer US sot FP 

11. Leaves typically 3-lobed or subcruci- 

form; taprooted perennials in soil. .. 
DE AE cod DOT TON 10. P. vaseyt. 
11. Leaves typically 3-lobed but not divided; 
woody-based perennials in rock crevices, 
or taprooted in soil ..... 18 P. parryi. 
10. Plants of Sonora and Sinaloa, Mexico, and 
Baja California Sur. ..... eoe 12. 
19. Plants subshrubs or suffrutescent peren- 
nidis. eur NEZX C S 13. 
13. Leaves densely puberulent and sub- 
canescent. ..... 22. P. leptoglossa. 
13. Leaves pilose and green. ...... 14. 
14. Leaves 3.5-11 em long; involu- 

eral bracts 10-13 mm long. 

EM _.. 21. P. cordifolia. 
14. Leaves 2.5-4.5 cm long; involu- 
eral bracts 5-7 mm long. .. . 
ee NICE 23. P. lloydii. 
12. Plants  herbaceous  perennials with 
rather thin, fleshy roots. ........- 15. 
15. Leaf blades 3-5 lobed, cleft, parted 
or rarely divided, the margins den- 
tate-lobed and acuminate; Baja 
California Sur. ..... 24. P. lobata. 
15. Leaf blades shallowly or strongly 
3.lobed, the margins serrate-den- 
tate; Sonora and Sinaloa, Mexico. 
Rl) MN OCA Um 25. P. palmeri. 

5. Ray corollas white, disc corollas yellow, except disc 

corollas cream-white in P. rosea and P. trichodonta. 
meds cp c a 16. 

Rhodora [Vol. 76 

16. Disc corollas eream-white. ... .. ....... 17. 
17. Leaves 1.8-3 cm long, 1-1.5 (2) em wide; 

ligules 2-2.5 mm long. ....... 15. P. rosei. 

17. Leaves 0.7-0.9 em long, 0.25-0.5 cm wide; 

ligules 1-1.5 mm long. .. 16. P. trichodonta. 

16. Disc corollas yellow. .................... 18. 



Achene margins thin, not calloused ; heads 
0.6-1 cm high, 0.6-1.5 cm wide, but possibly 
smaller in P. emoryi; Baja California Sur 
and islands, except P. emoryi widespread. 
DNUS 19. 
19. Ligules 6-10 mm long. ......... 20. 
20. Leaves usually thick and crisped, 
arachnoid-villous; Baja California 

Sur, coastal dune sand. [. 

. la. P. crassifolia var. crassifolia. 

20. Leaves usually rather thin and not 
crisped, densely short-pubescent to 
glabrous; Baja California Sur, in 

various soils. .................. 

. lb. P. crassifolia var. robusta. 

19. Ligules 1.5-4 (6) mm long (rarely ab- 
sent). ........ eee e eese. 21. 

2]. Plants  suffruticose perennials; 
Revillagigedo Islands. .......... 

ss 2. P. socorrensis. 

21. Plants delicate or robust annuals; 
widespread weed. .. 3. P. emoryi. 

Achene margins thin or prominently cal- 
loused; heads 3-7 mm high, 4-8 mm wide, 
but may be wider in P. turneri; Sierra 
Madre Occidental, Mexico, and foothills, 
and Arizona, except P. microglossa wide- 

spread. ........0.00.0.-0..0.000 004. 22. 
22. Achenes 1.8-3.5 mm long; pappus bris- 
tles 2 (3), 1.5-3 mm long. ....... 23. 

23. Leaves  deltoid-ovate to ovate- 
rhombic, the margins serrate, 



Perityle — Powell 247 

shallow-lobed, or serrate-crenate; 
central Arizona. ... 11. P. ciliata. 
Leaves ovate to subspathulate and 
entire to shallow-lobed, or ovate- 
cordate and serrate to serrate- 
lobed, or 2-3-pinnatifid to pedately 
divided; southern Arizona and 
Mexico. . ... SUM DNE 
24. Leaves entire, shallow-lobed 
or serrate; Durango, Mexico. 
ANES 13. P. hofmeisteria. 

24. Leaves pinnatifid or pedately 
divided with spathulate or 
linear segments; southern 
Arizona and adjacent Mexico. 

ow Haani P re er 25. 

25. Capitulescence of several 
heads clustered on short 
peduncles; achene mar- 

gins typically long-ciliate. 

_.. 12. P. coronopifolia. 

25. Capitulescence essentially 

of solitary heads; achene 

margins merely puberu- 

lent. 10. P. canescens. 

Achenes 1.3-2 mm long; pappus bris- 
tles 2 (or 0-2), 0.5-1.5 mm long. . 26. 



Leaves pinnately 3-5 divided, the 
divisions linear or nearly so. 
— D ! . 7. P. lineariloba. 

Leaves otherwise. ..... IDEE 

27. Heads 0.7-1.4 cm wide. .. 
MINNS 6. P. turneri. 

27. Heads 4-7 mm wide. . 28. 

28. Plants suffrutescent per- 
ennials; leaves densely 

. 8. P. microcephala. 

248 Rhodora [Vol. 76 

28. Plants delicate or robust 
annuals; leaves puberu- 
lent,  glandular-puberu- 
lent, or glabrous. 29. 
29. Ligules 1.5-3.5 mm 

long; upper pedun- 
cles usually copi- 
ously glandular-pu- 
berulent. .... 
.. 9a. P. microglossa 
var. microglossa. 

29. Ligules 3.5-4.5 mm 
long; upper pedun- 
cles usually sparsely 
to densely puberu- 
lent, rarely moder- 
ately  glandular-pu- 
berulent, . 
.. 9b. P. microglossa 
var. saxosa. 

1. Perityle crassifolia Brandeg. Proc. Calif. Acad. II, 3: 
147. 1891. TYPE: Mexico: BAJA CALIFORNIA: San Jose del 
Cabo, 6 Oct. 1890, T. S. Brandegee (Holotype, UC; isotypes, 
GH !, US(2) !). 

la. Perityle crassifolia var. crassifolia. 

Plants perennial, 10-75 cm high, younger plants with 
slender taproots, older plants with fleshy-woody taproots, 
younger plants erect and branching toward the top, older 
plants branching at the base, tending to be decumbent, 
stems pilose-villous to hirsute and glandular-pubescent; 
leaves mostly alternate, felty to touch, arachnoid-villous to 
short pilose-hirsute and glandular-pubescent, 2-3 (5) cm 
long, 1.5-2.0 (3.0) em wide, reniform to cordate in outline, 
thick and crisped, crenate to 3-5 lobed or dissected ; subses- 
sile to petiolate, the petioles 0.5-2.0 em long; capitulescence 
of 1-3 heads borne on peduncles 1-4 (6) em long; heads 
radiate, ca. 1.0 em high, 1.0-1.5 em wide, involucres hemi- 

1974] Perityle — Powell 249 

spherical; bracts numerous, ovate-lanceolate; ray flowers 
12-16, ligules white, 6-8 mm long often pinkish-tinged, ob- 
long to elliptic or subspathulate; dise flowers numerous, 
corollas yellow, 3.0-3.5 (4.0) mm long, throats tubular- 
funnelform; achenes 2.5-3.0 (4.0) mm long, lanceolate- 
obovate and curved, callous margins absent or very thin, the 
margins densely ciliate, ray achenes 3-angled and pubescent 
on faces, disc achenes 2-angled and glabrous on faces; 
pappus of a short crown of squamellae and 1 (rarely 2) 
slender, barbellate bristle, 2-3 mm long, the distal barbs 
retrorse; chromosome number, n = 19. 

Apparently restricted to deep coastal sand from Punta 
Arena to San Jose del Cabo, Cape Region of Baja California 
Sur, and Isla Coronados. Flowering year around. (Fig. 1). 


Mexico: BAJA CALIFORNIA SUR: Los Frailes, S of Cabo Pulmo, 
Arnaud (ps); San Jose del Cabo, Brandegee (DS, GH, NY); SW end 
of Isla Coronados, Carter 4274 (DS, SD, UC, US); Punta Frailes, Daw- 
son 1140 (us); 3 mi N of Los Frailes, Hastings and Turner 64-279 
(ARIZ) ; sand near shore, Isla Coronados, Moran 9121 (sp, vs); 0.5 
mi E of Eureka, Powell and Turner 1849 (SRSC, TEX) ; San Jose del 
Cabo, Purpus 444 (us) 274, 446 (vc); 0.6 mi S of Buena Vista, 
Wiggins 14747 (CAS, DS, GH, TEX). 

This taxon is best distinguished from var. robusta by 
its arachnoid-villous pubescence, distinct perennial habit 
with fleshy to woody taproots in older plants, and habitat 
in coastal sand. 

Some considerations suggest that var. crassifolia and var. 
robusta should be accorded specific rank. Plants from one 
collection of var. crassifolia (Powell and Turner 1849) and 
several collections of var. robusta have been grown from 
seed under identical greenhouse conditions. The character- 
istie morphologies of both taxa persisted under artificial 
conditions, but var. crassifolia flowered rarely while var. 
robusta flowered profusely. Vigorous intervarietal hybrids 
were obtained, but only two flowering heads were produced 
on only one of several plants. Meiosis was regular and 
pollen stainability was ca. 3%. The reproductive data are 

250 Rhodora [Vol. 76 

too meager to allow speculation about relationships. Also, 
I have seen specimens of var. robusta (among borrowed 
material) which approach var. crassifolia morphologically. 
I believe that it is best to retain varietal status for the taxa, 
following Everly (1947), pending populational study of the 
habit and habitat of var. crassifolia. 

1b. Perityle crassifolia var. robusta (Rydb.) Everly, Con- 
trib, Dudley Herb. 3: 382. 1947. 

Perityle robusta Rydb. N. Amer. Fl. 34: 16. 1914. TvPE: 
Mexico: BAJA CALIFORNIA: Cerralvo Island, 19 April 1911, 
J. N. Rose 16880 (Holotype, NY!; isotype, US!; isotype 
fragments UC (2) !). 

Perityle incompta Brandeg. Univ. Calif. Pub. Bot. 6: 
508. 1919. TYPE: Mexico: BAJA CALIFORNIA : Los Dolores, 
W. E. Bryant (uc!). 

Perityle macromeres Blake, Proc. Biol. Soc. Wash. 37: 
59. 1924. TYPE: Mexico: BAJA CALIFORNIA: La Paz 3 Feb 
1906, E. W. Nelson and E. A. Goldman 1483 (us!). 

Plants perennial or annual, with fleshy taproots to slen- 
der branching roots, erect with few branches to spreading 
in large clumps with many branches, stems short-hirsute to 
glabrous; leaves densely short-hirsute and glandular-pu- 
bescent to glabrous, 2-10 cm long, 1-7 cm wide, ovate to 
cordate in outline, rather thick and crisped to thin and not 
crisped, typically deeply 3-5 lobed with the lobes also in- 
dented or with somewhat irregularly dissected margins; 
heads 0.6-1.0 cm high, 0.6-1.5 em wide, involucres hemi- 
spherical to campanulate; ligules 6-10 mm long; disc corol- 
las 2-8 (4) mm long; pappus bristle usually 1, rarely 0-3; 
chromosome number, n = 19, 18. 

Growing in various soils, including those which are sandy 
or saline near the sea and occasionally among rocks, rather 
common in southern Baja California Sur and neighboring 
islands. Flowering year around. (Fig. 1). 


Mexico: BAJA CALIFORNIA SUR: Isla Espiritu Santo, Berry (CAS); 
Isla Magdalena, Brandegee (NY, UC, US); Isla Santa Margarita, 


Perityle — Powell 




Fig. 1. Distribution of P. crassifolia var. crassifolia (closed cir- 
cles); P. crassifolia var. robusta (open circles); P. aurea (closed 

squares); not plotted are P. socorrensis of the Revillagigedo Islands, 
and P. incana of Guadalupe Island. 


252 Rhodora [Vol. 76 

Brandegee (UC, US); San Jose del Cabo, Brandegee (POM); El 
Mogote Peninsula, La Paz Bay, Carter 2729 (CAS, DS, GH, UC, 
us); 45 km N of El Refugio, Carter, Alexander, and Kellogg 
2152 (DS, UC, US); Puerto Escondido, Carter and Kellogg 2875 (ARIZ, 
DS, GH, SD, UC, US); Isla Partida, Collins, Kearney, and Kempton 166 
(us); Isla San Francisco, Collins, Kearney, and Kempton 198 (vs); 
10 mi W of Comondu, Gentry 4083 (ARIZ, DS, GH, UC); dunes, San 
Nicholas Bay, Johnston 3720 (CAS, GH, UC, us); Isla Coronados, 
Johnston 3756 (CAS, DS, GH, NY, UC, US); dunes, Isla Monserrate, 
Johnston 3865 (CAS, GH, UC, US); beach, Agua Verde Bay, Johnston 
3893 (CAS, GH, NY, UC, US) ; dunes, Isla San Francisco, Johnston 3946 
(CAS, DS, GH, NY, UC, US) ; Isla Cerralvo, Johnston 4046 (CAS); Isla 
Espiritu Santo, Johnston 4081 (CAS, GH, NY, UC, US) ; Guadalupe Point, 
Concepcion Bay, Johnston 4150 (CAS, DS, NY, uc); E base of San 
Lazaro, Santa Maria Bay, Moram 3537 (ps, UC); Ensenada de los 
Muertos, Moran 3560 (ps, sD); S end of Isla Cerralvo, Moran 9616 
(ns, UC); Isla San Jose, Moran 3751 (ps, vC); Isla San Marcos, 
Moran 3948 (ps, UC) ; Isla Danzante, Moran 9209 (ps); W side of Isla 
Catalina, Moran 9329 (sp, UC); NE side of Isla San Jose, Moran 
9387 (sp); Isla Las Animas Rock, Moran 9433 (sp, vC, US); Isla 
San Diego, Moran 9593 (sp); Isla Carmen, Palmer 1 (GH, NY, US); 
10 mi N of Loreto, Powell and Sikes 1662 (SRSC, TEX); 2 mi NE of 
La Paz, Powell and Sikes 1681 (SRSC, TEX); 33 mi W of San Javier, 
Powell and Turner 1847 (sRSC, TEX); Isla Magdalena, Rose 16319 
(Ny, US); Isla Pichilinque, Rose 16529 (GH, NY, US); 8 mi W of 
San Miguel, Shreve 7125 (ARIZ, DS, GH, us); 17 mi N of La Paz, Sikes 
and Babcock 270 (SRSC, TEX); 2 mi S of Villa Constitucion, Sikes and 
Babcock 276 (SRSC, TEX); ca. 29 mi N of Villa Constitucion, Thomas 
8403 (CAS, DS, GH, US); 9 mi E of San Ignacio Wiggins 11360 (CAS, 
DS, GH, UC, US); San Gregoria, 12 mi NW of La Purisima, Wiggins 
11467 (ps, GH, UC); 0.6 mi S of Buena Vista, Wiggins 14748 (CAS, 
DS, TEX, UC); Salino Flat, near S end of Isla Espiritu Santo, Wiggins 
15595 (ps); Isla Partida, Wiggins 16160A (Ds). 

Perityle crassifolia var. robusta is more widespread than 
is var. crassifolia, and occupies a variety of edaphic condi- 
tions in coastal, insular, and inland areas in the southern 
portion of Baja California Sur. Two extreme forms of the 
morphologically variable var. robusta can be recognized. 
The plants of coastal habitats (e.g., near La Paz) usually 
are robust with fleshy, perennial taproots, while plants of 
the inland forms (e.g., near Commondu) are typically 
smaller with annual or weak perennial habits. Also, the 
inland form tends to be less pubescent and has thinner, less 

1974] Perityle — Powell 253 

crisped leaves than coastal plants. According to my inter- 
pretation, the “inland form” corresponds to P. incompta 
which Everly (1947) recognized as a distinct species. The 
“coastal form” conforms with Everly’s P. crassifolia var. 
robusta. Although the extremes of coastal and inland forms 
are evident, the existence of separate taxonomic entities 
has not been indicated. Instead, examination of exsiccata 
material and field studies have shown a rather complete 
morphological intergradation (e.g. Johnston 3946) between 
the two extremes. Hence P. incompta is treated here as 
synonymous with var. robusta. 

Further study is needed to clarify the adaptable habit of 
var. robusta, i.e., the perennial vs. annual conditions. Also, 
particular attention should be given to the habitats in which 
life forms occur. Those plants at La Paz grow in crusty 
saline soils, while plants of other populations may be in 
crevices of granite rocks, in rocky soil, or in fine sand. 

Another aspect of variability in var. robusta is presence 
or absence of pappus bristles, which seemingly is not taxo- 
nomically significant. As a generality, however, awnless 
forms occur on islands while awned forms are peninsular, 
but there are exceptions. 

2. Perityle socorrensis Rose, Bot. Gaz. 15: 118. 1890. 
TYPE: Mexico: Socorro Island, Revillagigedo Islands, Mar. 
1889, C. H. Townsend (Holotype, Us!; isotypes, GH !, NY !, 

Plants suffruticose perennials, 10-40 cm high, many 
branched and densely leafy; leaves typically alternate, 
densely short-hirsute and glandular-pubescent, semisuccu- 
lent, 2-6 cm long, 0.8-3.0 cm wide, ovate, deltoid-ovate, cor- 
date to subhastate, 3-5 shallow-lobed and serrate to crenate ; 
petioles 1.5-3.0 cm long; capitulescence of 1-2 (3-5) heads 
borne on peduncles 1.0-3.5 (6.0) cm long; heads typically 
radiate, rarely discoid, ca. 6 mm high, 7-10 mm wide, in- 
volucres campanulate to narrowly so; bracts lanceolate to 
oblanceolate or oblong-lanceolate; ray flowers ca. 10 or 
fewer, ligules white, 2-4 mm long, oblong to oblong-elliptic, 
rarely with a prominent inner lobelet; disc corollas yellow 

254 Rhodora [Vol. 76 

(?), 2.0-2.5 mm long, throats tubular-funnelform ; achenes mm long, oblanceolate-obiconical, oblong-oblanceo- 
late, to narrowly obconical typically curved, callous margins 
absent or very thin, the margins ciliate, ray achenes 
sparsely and minutely pubescent on faces, disc achenes 
glabrous or nearly so; pappus of a short to vestigial crown 
of squamellae and typically 2 (0-1) bristles, 1.5-2.0 mm 
long, the bristles retrorse barbellate (rarely lateral or an- 
trorse) distally; chromosome number, n = 19. 

Endemic to the Revillagigedo Islands, growing on sea 
cliffs and in soils near the shore. Flowering mostly in 
winter and spring. 


Mexico: REVILLAGIGEDO ISLANDS: Isla San Benedicto: Anthony 372 
(CAS, DS, GH, NY, UC, US); Barkelew 175 (ARIZ, DS, GH, NY, POM, UC, 
us); Mason 1684 (cas, GH, UC, US); Isla Clarion: Anthony 415 
(CAS, DS, POM, UC, US); Sulfur Bay, Howell 8347 (CAS, DS, NY, POM, 
us); W end of island, Mason 1578 (CAS, DS, GH, NY, POM, UC, US); 
Isla Socorro: Anthony 383 (ARIZ, CAS, DS, GH, POM, SD, UC, US); 
Barkelew 192 (ARIZ, DS, GH, NY, POM, UC, US); Academy Bay, Carl- 
quist 368 (CAS, RSA, UC); Brenner's Cove, Howell 8423 (CAS, DS, GH, 
uc, US); Graysons Cove, Moran 5922 (ARIZ, CAS, DS, GH, NY, RSA, SD, 
UC, US). 

Perityle socorrensis is a distinct species closely related to 
P. crassifolia from which it is distinguished by its typically 
2 pappus bristles, reduced squamellae, generally smaller 
floral features, short ligules (or absent), woody bases, leaf 
morphology, and its distribution. 

As discussed by Everly (1947), Perityle socorrensis is 
notably variable in ligule and pappus features. The ligules, 
typically short at 2-4 mm, are even shorter and aberrant- 
looking in many collections, while other specimens are 
without ligules. According to label data, discoid individuals 
may occur in populations with radiate forms, and thus no 
taxonomic significance is attributed to ligule variability. 
Typically, the number of pappus bristles per achene in P. 
socorrensis is two, but some individuals or populations may 
have only one bristle or none. Bristle number is correlated 

1974] Perityle — Powell 255 

with the island to island distribution: plants with two bris- 
tles on Socorro and San Benedicto Islands, and awnless 
forms on Clarion Island. No taxonomic significance is at- 
tached to bristle variability since exceptions are found on 
all the islands, and because other features are not corre- 
lated with the pappus differences. 

3. Perityle emoryi Torr. in Emory, Notes Mil. Rec. 142. 
1848. TYPE: California: mountains E of San Diego, 29 Nov. 
1846, Emory (NY!). 

Perityle nuda Torr. Pacif. R. Rep. 4: 100. 1857. TYPE: 
California: Williams’ River, 7 Feb. 1853-4, J. M. Bigelow 

Perityle emoryi var. nuda A. Gray, Bot. Calif. 1: 397. 

Perityle emoryi S. Wats. Proc. Amer. Acad. 11: 116. 1876. 
Not P. emoryi Torr. 1848. 

Perityle californica A. Gray, Syn. Fl. N. Amer. 1: 321. 
1884. Not P. californica Benth. 1844. 

Perityle californica var. nuda A. Gray, Syn. Fl. N. Amer. 
1: 321. 1884. 

Perityle fitchii var. palmeri A. Gray, Syn. Fl. N. Amer. 1: 
321. 1884. 

Perityle fitehii Green, Bull. Calif. Acad. 2: 403. 1887. 
Not P. fitchii Torr. 1857. 

Perityle californica Vasey, Proc. U, S. Nat. Mus. 11: 368. 
1889. Not P. californica Benth. 1844. 

Perityle rothrockii Rose, Bot. Gaz. 15: 114. 1890. TYPE: 
Nevada: 1872, Wheeler (us!). 

Perityle greenei Rose, Bot. Gaz. 15: 117. 1890. TYPE: 
California: Santa Cruz Isl, July-Aug. 1886, E. L. Green 
(Holotype, ND?; isotypes, DS!, NY!, UC!, US!). 

Perityle emoryi var. orcuttii Rose, Bot. Gaz. 15: 117. 
1890. TYPE: Mexico: BAJA CALIFORNIA. Canyon Cambel- 
los(?) July, 1884, C. R. Orcutt (Holotype, US!; isotype 

Perityle grayi Rose, Bot, Gaz. 15: 118. 1890. TYPE: 
Mexico: Guadalupe Isl., 1875, E. Palmer 44 (Holotype, Us?; 
isotypes, GH !, NY!). 


256 Rhodora [Vol. 7 


Laphamia nuda Benth & Hook. ex Jacks. Ind. Kew. 
30. 1895. 

Laphamia emoryi Benth & Hook, ex J acks. Ind. Kew. 2: 
30. 1895. Partial Synonomy of South American P. emoryi 
(= Closia). 

Closia elata Phil Fl. Atac. 31 and Viage Des. Atac. 19, 
205. 1860. Type not examined. 

Closia discoidea Phil. Fl. Atac. 31 and Viage Des. Atac. 
205. 1860. Type not examined. 

Perityle emoryi Torr. var. elata. (Phil.) I. M. Johnston, 
Contr. Gray Herb. 85: 127. 1929. 

Perityle discoidea (Phil) I. M. Johnston, Contr. Gray 
Herb. 85: 128. 1929. 

Plants delicate or robust annuals, 2-60 cm high, usually 
herbaceous or the lower stems woody, erect or spreading 
with few to many stems, puberulent to hirsute and glandu- 
lar-pubescent; leaves mostly alternate, hirsute to glandular- 
pubescent, 2-10 cm long 1-5 cm wide, ovate, cordate, sub- 
orbieular, to triangular in outline, the margins deeply 
toothed, lobed, cleft, or divided, with the segments also 
indented to irregularly dissected; petioles 0.3-4.0 cm long ; 
capitulescence of 1 to many heads borne on peduncles 0.1- 
7.0 em long ; heads radiate (rarely discoid), 0.4-1.0 cm high, 
0.4-1.0 em wide; involucres hemispherical or campanulate; 
bracts numerous, lanceolate, oblanceolate, to ovate-lanceo- 
late; ray flowers 8-12 (14), ligules white, usually oblong, 
1.5-4.0 (6.0) mm long, 1.5-3.0 mm wide, rarely rudiment- 
ary; disc flowers numerous, corollas yellow, 2.0-2.5 (3.8) 
mm long, throats tubular to tubular-funnelform; achenes 
(1.5) 2-3 mm long, suboblong, oblanceolate, or subcuneate, 
the outer ones often curved, margins thin, not calloused, 
the margins long- or short-pubescent (ciliate), outer 
achenes 2-3 (4) angled and often pubescent on the faces, 
inner achenes 2-angled with glabrous (rarely puberulent) 
faces; pappus of a vestigial or conspicuous crown of squa- 
mellae and 1 slender bristle, (0.8) 1.0-2.5 (3.5) mm long, 
antrorse-, lateral-, or retrorse-barbed especially at the tip. 

1974] Perityle — Powell 257 

Fig. 2. Generalized distribution of P. emoryt (stippled); South 
American distribution in Chile and Peru not plotted. 

258 Rhodora [Vol. 76 

or the bristle absent; chromosome number, tetraploid (n = 
32-36) or hexaploid (n = 50-58). 

Widespread weed of desert southwestern United States, 
Sonora, Mexico, Baja Californian peninsula, and neighbor- 
ing islands; continental disjunct in Chile and Peru. Fower- 
ing mostly in winter and spring, but also year around, de- 
pending upon latitude. (Fig. 2). 


MEXICO: Baja California: Ensenada, Anthony 180 (CAS, Ds, UC, 
us); 8.2 mi S of Socorro, Constance 3119 (DS, GH, LL, UC, US); 
Bahia de los Angeles, Cowan 2265 (TEX); 14 mi S of Santa Rosa- 
lillita, Powell and Sikes 1647 (SRSC, TEX); 41.6 mi S of Mexicali, 
Powell and Turner 1710 (SRSC, TEX); Arroyo Calmalli, Purpus 23 
(CAS, NY, POM); 14 km NW of Colonia Guerrero, Raven, Lewis, and 
Thompson 12193 (GH); 8.5 km E of El Rosario, Raven, Mathias, and 
Turner 12433 (ARIZ, GH); 15.9 mi N of San Felipe, Raven 14775 
(ps, uc); N side of El Arco, Sikes and Babcock 295 (SRSC, TEX); 
2 mi N of Miission de San Borja, Wiggins and Wiggins 14851 (DS); 
9 mi S of Puertocito, Wiggins and Wiggins 15877 (Ds, US). Baja 
California Sur: Sierra de la Trinidad, Cape Region, Brandegee (UC, 
US); near Santa Rosalia, Ferris 8702 (Ds); Santo Domingo, Gander 
9792 (sp); 10 mi W of Comondu, Gentry 4083 (ARIZ, UC); San 
Francisquito Bay, Johnston 3566 (CAS, GH, US); Concepcion Bay, 
Johnston 4150 (GH, US); E base of San Lazaro, Santa Maria Bay, 
Moran 3537 (sp); 18.6 mi. S of Mulege, Powell and Turner 1836 
(SRSC, TEX); ca. 20 mi N of Santa Rosalia, Sikes and Babcock 285 
(SRSC, TEX); 8 mi NW of San Ignacio, Sikes and Babcock 288 (SRSC, 
TEX); San Ignacio, Wiggins 16233 (ps); 3 mi N of El Barril, Wig- 
gins 16864 (ns); Mulege, Wiggins and Wiggins 18070 (us); 4 mi 
S of El Arco, Wiggins and Wiggins 18191 (CAS, DS). Islands: ISLA 
ANGEL DE LA GAURDA: Wiggins 17010 (ps). ISLA CARMEN: Moran 
9181 (sp). ISLA CEDROS: Anthony 283 (CAS, DS, GH, NY, POM, UC, US); 
Haines and Hale (CAS, GH, LL, NY, SD, UC, US); Howell 10684 (CAS, 
DS, GH, NY, POM, US). ISLA CORONADOS: Moran 6556 (sD); Palmer 16 
(GH, NY, US). ISLA GUADALUPE: Anthony 233 (CAS, DS, GH, UC, US); 
Carlquist 489 (RSA); Mason 1502 (CAs, GH, US); Moran 2900 (DS, 
GH), 5628 (DS, RSA, SD, UC); Palmer 891 (Ny, UC, US); Rose 16003 
(GH, NY, US). ISLA ILDEFONSO: Moran 9062 (sp); Wiggins 18254 
(CAS, DS). ISLA NATIVIDAD: Brandegee (UC); Moran 10797 (sp). 
ISLA PARTIDA: Johnston 3235 (CAS, US); Wiggins 17268 (DS). ISLA 
BENITO: Anthony 271 (DS, GH, POM, UC); Palmer 914 (ARIZ, CAS, 

1974] Perityle — Powell 259 

DS, NY, US). ISLA SAN ESTEBAN: Wiggins 17218 (DS). ISLA SAN 
LORENZO: Moran 8895 (SD). ISLA SAN MARCOS: Johnston 3622 (CAS, 
GH, UC, US). ISLA SAN PEDRO MARTIR: Johnston 3148 (CAS, GH, UC, 
us); Moran 8810 (SD). ISLA TIBURON: Johnston 3268 (CAS, us). 
ISLA TODOS SANTOS: Moran 16214 (sD). ISLA TORTUGA: Wiggins 
17367 (ps). Sonora: Puerto Penasco, Breedlove 1397 (ns); 27.4 mi 
S of Sonoyta, Breedlove 1389 (Ds, TEX); vicinity of Libertad, Graham 
3803 (ps); 3 mi W of Kino, Heckard 1575 (JEPS); 1 mi E of San 
Carlos Resort, Powell and Sikes 1685 (SRSC, TEX); Empalme, Rose, 
Standley, and Russell 12636 (GH, NY, us); 5 mi NW of Caborea, 
Shreve 1532 (ARIZ); 4 mi NW of Caborea, Wiggins 8233 (DS, GH, 
UC, US). 

U.S.A.: Arizona: COCONINO CO.: Havasu Canyon, Grand Canyon, 
Howell 26537 (ARIZ, CAS, SMU); GILA CO.: Roosevelt Dam, Eastwood 
6247A (CAS). MARICOPA CO.: Phoenix, Eastwood 6159 (CAS); near 
Tempe, Gillespie 8808 (DS, GH, POM, UC, us); 2 mi NW of Scottsdale, 
Russell 10911 (SMU, UC). MOHAVE CO.: 7 mi NW of Alamo, Benson 
10074 (pom); 3.4 mi SE of Boulder Dam, Carter and Chisaki 3230 
(ARIZ, DS, LL, NY, RSA, SMU, UC, US) ; Toroweap, Grand Canyon, Cottam 
13001 (UT); 62 mi N of Wickenburg, Sikes 106 (SRSC). PIMA CO.: 
10 mi W of Bates Well, Growler Mts., Benson 9926 (PoM); Gunsight 
Peak, Fosberg 7865 (POM); ca. 5 mi N of Ajo, Gould and. Macbride 
4130 (ARIZ, GH, NY, UC); Alamo Canyon, Ajo Mts, Huey 24351 
(cH, sp); Dripping Spring, Organ Cactus Natl. Mon. McClintock 
52-37 (CAS). PINAL CO.: Sacaton, Gilman 350 (ARIZ); 10 mi W of 
Maricopa, Russell 11348 (smu). YUMA CO.: Gila Mts., near US 80, 
Barr and Lange 64-174 (ARIZ); S end of Castle Dome Mts., Parker, 
Wright and Lowe 7789 (ARIZ, DS, NY, RSA, us); 15.6 mi E of Yuma, 
Powell and Turner 1704 (SESC, TEX); S end of Cunningham Pass, 
Harcuvar Mts., Wiggins 8452 (ps, GH, UC, us). California: IMPERIAL 
co.: 8 mi from Niland to Blythe, Balls 12921 (RSA); 7 mi N of 
All American Canal spillway, Wiggins 8613 (DS, NY, POM, UC, US). 
Inyo co.: Furnace Creek, Death Valley, Carpenter (JEPS) ; Funeral 
Mts., Coville and Funston 324 (us); Hanaupah Canyon, Coville and 
Gilman 392 (vs); Darwin Falls, Hitchcock 6219 (uc); Surprise 
Canyon, Howell 3964 (CAS); Panamint Valley Smith 86 (JEPS). 
KERN CO.: Last Chance Canyon, El Paso Range, Twisselman 11863 
(JEPS). LOS ANGELES CO.: Santa Catalina Isl., Brandegee (UC); 
Santa Monica Mts., Epling (DS, NY, RSA); Anacapa Isl., Howell 3816 
(JEPS); San Clemente Isl., Raven 17345 (RSA, UC). RIVERSIDE CO.: 
Palm Springs, Abrams 11012 (ps); San Jacinto Range, Benson 4167 
(pom): 8 mi NE of Desert Center, Wiggins 9675 (DS, GH. RSA, UC). 
SAN BERNARDINO CO.: 29 Palms, Alexander and Kellogg 870 (UC); 
near Parker Dam, Brenckle 51140 (SMU, uc); 89 mi N of Needles, 
Ferris 7224 (ps); between Kelso and Baker, Jepson 20590 (JEPS). 
SAN DIEGO CO.: 23.1 mi NW of Coyote Wells, Breedlove 1856 (DS); 

260 Rhodora [Vol. 76 

Yaqui Wells, Eastwood 2644 (CAS, GH, UC, US); Jacumba, Nelson 
11182A (DS, GH, NY, POM, UC); Borego Park, Wolf 8462 (ARIZ, au, 
NY, RSA). SANTA BARBARA CO.: Santa Cruz Isl., Balls and Blakley 
23727 (RSA, UC). VENTURA CO.: Point Mugu, Howell 3733 (cas). 
Nevada: CLARK COo.: between Las Vegas and Boulder Beach, Cron- 
quist 9844 (NY, UC); 1 mi below Boulder Dam, Grater 51 (vc); 8 mi 
SW of Davis Dam, Gullion 258 (Uc). 

CHILE: Atacama: Chanaral, Beetle 26164 (GH, UC); vicinity of 
Caldera, Gigoux (GH); below Agua EI Huerto, Johnston 3678 (vs); 
vicinity of Potrerillos, Johnston 4740 (GH, US); vicinity of Puerto 
de Chanaral, Johnston 4794 (GH, US); vicinity of Copiapo, Johnston 
5028 (GH); Caldera, Johnston 5056 (GH); vicinity of Caleta Pan de 
Azucar, Johnston 5829 (Us); Vallenar, Werdermann 160 (GH); 
Tierra Amarilla, Werdermann 406 (GH, NY); Quebrada Paipote, 
Werdermann 448 (GH, NY). Antofagasta: Taltal, Jaffuel 984 (GH): 
Tocopilla, Jaffuel 1014 (GH); Antofagasta, Jaffuel 1126 (GH); 6 km 
N of Puerto Tocopilla, Johnston 3585 (Us). 

ECUADOR: Galapagos Isles: Cerros Isl, Stewart 40 (cas). 

PERU: Mts. near Chosica, Lima-Oroya Railroad, Weberbauer 
5820 (GH, US). 

The morphological variation of Perityle emoryi is at- 
tested to by its considerable synonymy. This widespread 
annual, polyploid weed exhibits variation in nearly all as- 
pects of plant form. I have carefully examined the bulky 
exsiccatae available to me, giving particular attention to 
plant size, leaf morphology, head size, presence or absence 
of ligules and pappus bristles, and geographic distributions. 
None of the variable morphological aspects appear to have 
populational significance, and thus, in my judgement, do 
not require taxonomic recognition. I do note, however, that 
several atypical collections of P. emoryi from Magdalena 
Island resemble P. crassifolia and Amauria brandegeana, 
and thus the desirability of further study of Perityle from 
this locality is indicated. 

Perityle emoryi is related to and probably derived from 
the diploid P. crassifolia var. robusta, from which it is 
delimited by habit, smaller heads, shorter ligules, usually 
Shorter disc corollas, and chromosome number. With poorly 
preserved specimens it is often difficult to distinguish these 
taxa, but I have not had difficulty recognizing them in the 
field. Perityle emoryi is also remarkably similar to Amauria 

1974] Perityle — Powell 261 

brandegeana in superficial morphology. A discussion of the 
later similarity and distinguishing traits is to be found 
elsewhere (Powell, 1972a). 

It is assumed that P. emoryi achieved wide distribution 
as a result of the increased adaptability provided through 
a combination of polyploidy, reproductive self-compatibility 
and annual habit. That polyploids often exceed their di- 
ploids in distribution is well-known (Stebbins, 1950), and 
the advantage of self-compatibility in the establishment of 
disjunct colonies is clear. Indeed, Raven (1963) has sug- 
gested that P. emoryi might have attained bicontinental 
distribution as a result of long-distance dispersal from the 
Sonoran Desert, and he indicated the advantage of self- 
compatibility to such long-distance dispersal. More spe- 
cifically, I believe that the South American P. emoryt orig- 
inated from the vicinity of southern Baja California. It is 
in southern Baja California that the ancestors of P. emoryt 
are found. Furthermore, geographic variation in length of 
pollen spines (the spines ornamenting pollen walls) sug- 
gests that South American and Mexican P. emoryi had 
common areal origin. In measuring the pollen spines of 
some 20 populations of P. emoryi, it was found that the 
spine length of South American populations averaged the 
same or slightly longer than that of Mexican populations, 
while the spine length of United States populations was 
nearly twice as short as the others (Powell & Miller, un- 
published). From this information it can be postulated 
that P. emoryi originated and became established in Baja 
California, migrated to South America by long-distance 
dispersal in late Pliocene or Pleistocene (Raven, 1963), and 
subsequently advanced to the north, occupying desert areas 
in southwestern United States. 

Further study of the South American Perityle emoryi is 
needed, particularly chromosomal analyses and living plant 
comparisons. At least two forms can be recognized among 
the South American specimens I have seen, and this was 
discussed to some extent by Johnston (1929). One form 
(= P. discoidea (Phil.) I. M. Johnst.) deviates from North 

262 Rhodora [Vol. 76 

American P. emoryi in smaller habit, numerous, smaller 
and tightly clustered heads with no conspicuous ligules, and 
a pappus bristle on the achenes. Another form (= P. 
emoryi var. elata (Phil.) I. M. J ohnst.) is variable in habit, 
has large and small heads, longer and short ligules, and 
awnless achenes. I have not been able to make taxonomic 
distinctions between the South American and North Amer- 
ican forms, pending further study, and thus have placed 
Johnston’s combinations in Synonymy. It should be noted 
that only a partial synonymy of South American P. emoryi 
(Closia, in part?) is included since I have not seen speci- 
mens or types upon which several other Closia names are 
based. A few other collections from South America closely 
resemble P. crassifolia (e.g., Johnston 5023) or Amauria 
brandegeana (e.g., Werdermann 160; Morong 93). As 
mentioned above, these similarities are also seen in a few 
Mexican specimens. But it is also possible that both P. 
crassifolia and Amauria brandegeana once made their way 
to South America even though they have not become estab- 
lished there. 

4. Perityle aurea Rose, Contr. U. S. Nat. Herb. 1: 84. 1890. 
TYPE: Mexico: BAJA CALIFORNIA: Santa Rosalia, 24 Feb.- 
3 Mar. 1889, E. Palmer 185a (Holotype, Us!; isotypes, CAS !, 
GH !, NY!). 

Plants annual, 10-60 cm high, erect or spreading, stems 
of larger plants succulent, reddish, nearly glabrous; leaves 
alternate, puberulent to glabrous, slightly thick in texture, 
1.7-9.0 em long, 1-4 cm wide, subreniform to ovate in out- 
line, basically 3-lobed with dentate-serrate lobes and mar- 
gins; petioles 0.6-5.0 em long; capitulescence of 1-3 heads 
borne on rather short peduncles 1-2 (3) em long; heads 
radiate, 5-8 mm high, 5-8 mm wide, involucres campanulate 
to narrowly so; bracts broadly lanceolate; ray flowers ca. 
10, ligules yellow, oblong-elliptie to obovate; dise corollas 
yellow, 2-3 mm long, throats tubular to funnelform; achenes 
2-3 mm long, narrowly oblanceolate to suboblong, with thin 
callous margins, the margins densely white-ciliate, ray 
achenes pubescent on faces, disc achenes glabrous or nearly 

1974] Perityle — Powell 263 

so on faces, pappus of a conspicuous crown of squamellae 
and 1 bristle (rarely 0) 1.5-2.5 mm long, the bristle retrorse 
barbelate with few barbs at tip; chromosome number, 
n= M. 

Restricted in soils and among rocks, near Santa Rosalia 
of Baja California Sur and a few islands in the Sea of 
Cortez. Flowering Jan-April. (Fig. i9» 


Mexico: BAJA CALIFORNIA: summit and N slope of high peak near 
SE corner of Isla San Esteban, Moran 8845 (sD, US); summit of NE 
peak, Isla San Esteban, Moran 13044 (sb); arroyo on S Isla San 
Lorenzo, Moran 13060 (SD). BAJA CALIFORNIA SUR: Isla San Marcos, 
Johnston 3614 (CAS, GH, UC, US); Isla San Marcos, Moran 3965 
(ps); Santa Rosalia, Palmer 185a, 185 (CAS, GH, us); 8 mi W of 
Santa Rosalia, Powell and Turner 1826, 1829 (SRSC, TEX); 9 mi N 
of Santa Rosalia, Powell and Twrner 22219 (SRSC, TEX); 10 mi N 
of Santa Rosalia, Reed 6225 (ps, POM) ; ca 12 mi N of Santa Rosalia, 
Sikes and Babcock 281 (SRSC, TEX). 

Two growth forms of Perityle aurea are evident in the 
field, and both have been preserved in existing collections 
of the species. One form is of small plants, superficially 
resembling the Baja California populations of P. californ- 
ica, and the other form is of taller, more robust and succu- 
lent plants resembling P. crassifolia. The small form is 
known to occur in roadside soils where dry conditions pre- 
vail, and the large form occurs in places that receive and 
retain more water. The growth forms likely are ecologically 
controlled. I have seen both forms growing only a few 
paces apart, and when seeds of both plant sizes were grown 
under identical greenhouse conditions, only the larger, more 
succulent forms developed. The island specimens of P. 
aurea differ slightly from the mainland forms, most notably 
in achene morphology and in being awnless, although awned 
forms also occur on San Esteban Island. 

Perityle aurea seemingly is related to P. crassifolia, from 
which it is readily delimited by yellow ligules, chromosome 
number (n = 17), and distribution. An affinity of P. aurea 
with the P. crassifolia alliance is not indicated by its yellow 

264 Rhodora [Vol. 76 

ligules and chromosome number. All other members of the 
P. crassifolia group have white ligules and have x — 19. 
My first impression from superficial morphological exam- 
inations was that the species was related to P. californica 
(n = 13, 12, 11) and P. cuneata (n = 16, 12), both annual 
yellow-rayed taxa with habits similar to the small growth 
form of P. aurea. But P. aurea exhibits a combination of 
vegetative and floral features which characterize only the 
P. crassifolia alliance: robust and rather succulent habit, 
achenes with callous margins absent or thin, other unique 
achene morphology, and pappus bristles retrorsely barbed 
at the tips. Also, chromatographic studies have shown that 
P. aurea, like members of the P. crassifolia group, pro- 
duces the abundant yellow flavonoid compounds that have 
been detected elsewhere in sect. PERITYLE only in the P. 
parryi group of Texas. I conclude that P. aurea evolved, 
probably from P. crassifolia var. robusta, through specia- 
tion that involved adaptation of the annual habit and aneu- 
ploid reduction in chromosome number. Artifieial hybrid- 
izations have strongly suggested that the above taxa are 
reproductively isolated (Powell, 1972c). 

5. Perityle incana Gray, Proc. Amer, Acad. 11: 78. 1876. 
TYPE: Mexico: BAJA CALIFORNIA: Guadalupe Island, Feb.- 
May, 1875, E. Palmer 43 (Holotype, GH!; isotypes, NY, 

Nesothamnus incanus (Gray) Rydb. N. Amer. Fl. 34: 12. 

Plants shrubby, 10-40 (80) em high, many branched and 
densely leafy, stems tomentulose-canescent; leaves alter- 
nate, thickish, tomentulose-canescent, 6-14 em long, 4-7 em 
wide, deeply 3-divided or palmate, the divisions again 
deeply lobed or cleft; petioles 1.5-7.0 em long; capitules- 
cence of many tightly clustered heads (naked corymbs) 
borne on short peduncles; heads discoid, 5-7 mm high, 4-8 
mm wide, involucres campanulate; bracts broadly oblong- 
lanceolate to broadly linear; disc corollas yellow, 2-3 mm 
long, throats narrowly campanulate or broadly tubular; 

1974] Perityle — Powell 265 

achenes 2.0-3.8 mm long, narrowly obdeltoid to narrowly 
oblanceolate, with prominent callous margins, the margins 
coarse-ciliate, the faces coarse-pubescent ; pappus crown of 
2 broad, laciniate squamellae, 0.5-0.9 mm long, and rarely a 
narrow, fimbriate scale, 1.0-1.2 mm long; chromosome num- 
ber, n = ca. 50-57. 

Endemic to Guadalupe Island and islets, growing on cliffs 
near the sea. Flowering winter, spring, and early summer. 


MEXICO: Baja California: ISLA GUADALUPE: Anthony 249 (DS); 
Carlquist 440 (RSA, uc); Barracks Cove, Copp 157 (DS) ; Franceschi 
7 (A, DS, POM, UC); Howell 8172 (CAS); Outer Islet, Lindsay 2621 
(sp); Islote Negro, Mason 1519 (ARIZ, CAS, DS, us); Moran 2904 
(ps, RSA); Outer Islet, Moran 5682 (CAS, DS, NY, SD, uc); North 
Twin Canyon, Moran 6133 (sp); N end of island, Moran 6435 (DS, 
sp); cliffs of the Lower Circus, Moran 12023 (sp); N side of Mt. 
Augusta, Moran 12042 (DS, SD, uc); SW Oak Canyon, Moran 13792 

This cliff-dwelling Guadalupe Island endemic is perhaps 
the most distinct species of sect. PERITYLE. Its identifying 
features include: shrubby habit; tomentulose-canescent 
pubescence ; clustered, naked capitulescence; coarse pubes- 
cenct achenes; pappus of broad, laciniate squamellae and 
rarely à narrow, fimbriate scale. Considering its relatively 
large, shrubby habit, unique character, and isolated distri- 
bution, Perityle incana would seem to be a primitive mem- 
ber of the genus, although this possibility is not strength- 
ened by its hexaploid chromosome number. According to 
Reid Moran (personal communication), the taxon is not in 
danger of extinction, largely because its habitat is out of 
reach of the goats which have overrun the island for more 
than a century. 

It is clear that Perityle incana does not belong with any 
of the related-species groups, and thus stands as an anoma- 
lous species. In fact, the species could be recognized as a 
montotypic genus with little change in the taxonomic nat- 
uralness of the subtribe. I suspect, however, that the taxon 
is distantly related to the P. crassifolia group and is best 

266 Rhodora [Vol. 76 

treated as a member of Sect. PERITYLE. Perityle incana is 
similar to Pericome in habit and capitulescence characters, 
and perhaps shares ancestral affinity with that genus. 

6. Perityle turneri Powell, Madrofio 21: 456-457. 1972. 
TYPE: Mexico: DURANGO: 3.4 mi E of Ey Palmito, 2 Apr. 
1970, A. M. Powell 1858 (Holotype, TEX ! ; isotype, SRSC!), 

Plants herbaceous to suffrutescent perennials, decumbent, 
prostrate, or semierect, stems Spreading 20-45 em long, 
often purplish, lower stems often rooting at the nodes, 
upper stems puberulent to densely so, short-pilose, or sub- 
tomentose; leaves mostly opposite, puberulent to densely so, 
short-pilose to densely so, or subtomentose, often purplish, 
3.0-4.5 (8.5) em long, 1.5-3.0 (4.0) em wide, ovate, deltoid, 
or subcordate, the apexes acute or attenuate, the Margins 
serrate, serrate-crenate, Serrate-lobed to doubly so, the ser- 
rations or lobes acute or acuminate; petioles 1.0-1.5 (3.0) 
cm long; capitulescence of 1-3 (4) heads on peduncles (1.0) 
2-6 cm long; heads radiate, subglobose, 5-7 mm high, 0.7-1.4 
cm wide, involucres hemispherical to broadly campanulate; 
receptacles conical; ray flowers ca. 12-18, ligules white, (3) 
6-9 mm long, oblong; disc corollas yellow, (1.8) 2.0-2.8 
(3.0) mm long, throats short-campanulate, short-funnel- 
form, rarely narrowly tubular-funnelform ; achenes 1.5-2.0 
mm long, obovate to oblong-ovate, with thin or prominent 
callous margins, the margins ciliate, the faces puberulent in 
the center; pappus of a prominent crown of squamellae and 
2 (rarely 1) unequal bristles, the longest 0.6-1.5 mm long; 
chromosome numbers n — 17,n — 17 II 4 I. 

Moist seeps and canyons, higher elevations to ca. 9500 
feet, southern Chihuahua and Durango in the Sierra Madre 
Occidental. Flowering spring and fall. (Fig. 3). 


Mexico: CHIHUAHUA, 4 mi SW of Villa Matamoros, Correll and 
Gentry 22819 (LL); near La Rocha, NE slope of Sierra Mohinora, 
Correll and Gentry 23109 (LL); Burro Canyon near Parral, Pringle 
18650 (ARIZ, Cas, GH, SMU, TEX, UC, US). Durango. Quebrada San 
Juan, ca. 50 mi W of Durango and 23 mi NW of Los Coyctes RR, 

1974] Perityle — Powell 267 

Fig. 3. Distribution of P. turneri (closed squares) ; P. lineariloba 
(closed star); P. microcephala (closed triangles) ; P. microglossa var. 
microglossa (closed circles); P. microglossa var. saxosa (open tri- 
angles); P. ciliata (open squares); P. coronopifolia (open circles) ; 
P. hofmeisteria (open star); not plotted is P. canescens. 

268 Rhodora [Vol. 76 

Cronquist 9568 (NY, SMU, TEX, US); 2 km S of El Espinozo del 
Diablo, and 65 km SW of El Salto, Cronquist and Fay 10786 (NY); 
6 mi W of La Ciudad, Flyr 273 (TEX); ca. 35 mi W of El Salto, 
Gentry and Arguelles 18210 (Us) ; San Ramon, Palmer 60 (GH, NY, 
UC, US); 2.2 mi E of El Palmito, Powell and Turner 1857 (SRSC, 
TEX); 12 mi W of La Ciudad, Sikes and Babcock 380 (SRSC, TEX). 
SINALOA. 0.9 mi W of El Palmita, ca. 47 mi E of Concordia, Breed- 
love 1721 (Ds). 

Plants of this remarkable species were rarely collected 
until part of its high-altitude distributional range was made 
accessible by the road cut across the Sierra Madre Occiden- 
tal from Durango to Mazatlan. At the type locality near 
Mex. 40, Perityle turneri is abundant in wet mats formed 
by dense growths of Selaginella and small ferns. In overall 
aspect, at least at the type locality, the taxon shows striking 
resemblance to members of the tribe Astereae. 

Perityle turneri is related to P. lineariloba, from which 
it is easily delimited by leaf shape, and to P. microcephala, 
from which it can be distinguished by its capitulescence and 
head size. Further discussion concerning the taxonomy of 
this species is available elsewhere (Powell, 1972e). 

7. Perityle lineariloba Rydb. N. Amer. Fl. 34: 18. 1914. 
TYPE: Mexico: DURANGO: San Ramon, 21 Apr.-18 May 1906, 
E. Palmer 89 (Holotype, NY!; isotypes, GH!, UC!, US!). 
Plants seemingly perennial, with semi-annual roots, stems 
lignescent near the base, decumbent, ca. 30 cm long, grayish- 
puberulent above; leaves mostly opposite, finely puberulent, 
2.5-6.0 em long, 1.5-3.5 cm wide, pinnately 3-5 divided, the 
divisions linear or nearly so and entire or cleft into linear 
segments; petioles 0.7-2.0 em long; capitulescence of solitary 
heads on long peduncles, 4-10 cm long; heads radiate, 5-6 
mm high, 5-7 mm wide, involucres broadly campanulate; 
ray flowers ca. 10-15, ligules white, 4-7 mm long, oblong ; 
disc corollas yellow (?), 1.8-2.0 mm long, throats narrowly 
campanulate-funnelform; achenes 1.3-1.5 mm long, nar- 
rowly obovate to oblong-obovate, with thin or prominent 
callous margins, the margins and faces short-pubescent ; 
pappus of a conspicuous crown of squamellae and 1 (rarely 

1974] Perityle — Powell 269 

none) delicate bristle, ca. 0.5 mm long; chromosome num- 
ber, unknown. 

Known only from the type collection. (Fig. 3). 

The distinguishing characteristics of Perityle lineariloba 
include its pinnately divided leaves, long peduncles, and rel- 
atively large heads with long ligules. Most of its floral 
features are like those of P. turneri, to which the species is 

8. Perityle microcephala A, Gray, Proc. Amer. Acad. 21: 
391. 1886, TYPE: Mexico: CHIHUAHUA: rocky hills near 
Chihuahua, 8 Oct. 1885, C. G. Pringle 571 (Lectotype, GH ! ; 
isolectotype, US!). LECTOPARATYPE: Chihuahua: mountains 
above Batopilas, Oct, 1885, E. Palmer 268 (GH) ; isolecto- 
paratypes, NY! US !). 

Plants suffrutescent perennials, 20-40 cm high, many- 
stemmed and densely leafy, erect or decumbent, densely 
grayish-puberulent to pilosulous; leaves mostly opposite, 
densely grayish-puberulent, 1.5-3.5 (5.0) em long, 0.8-2.0 
(3.0) em wide, deltoid-ovate, ovate, to subcordate, the mar- 
gins serrate-crenate to doubly so, or shallow-lobed; petioles 
0.3-1.0 (1.8) cm long; capitulescence of severa] heads tight- 
ly clustered on short peduncles; heads radiate, 3-4 mm high, 
4-5 mm wide, involucres short-campanulate; ray flowers 
ca. 8-12, ligules white, 2-5 mm long, oblong; disc corollas 
yellow, often purple-tinged, 1.5-2.0 mm long, throats nar- 
rowly campanulate-funnelform; achenes 1.5-1.8 mm long, 
narrowly obovate to oblong-ovate, with callous margins, the 
margins and faces short-pubescent; pappus of a prominent 
crown of squamellae and 2 (rarely 1) unequal, delicate 
bristles, the longest 0.5-1.0 mm long; chromosome number, 

Sierra Madre Occidental of Chihuahua and Sonora. 
Flowering Aug-Nov, and probably in the spring also. 
(Fig. 3). 

Mexico: CHIHUAHUA. Damon s.n. (UC); Guicorichi, Rio Mayo, 

2'(0 Rhodora [Vol. 76 

Gentry 1973 (ARIZ, UC, US); near Batopilas, Goldman 215 (GH, NY, 
us); Madrono Mine, ca. 5 km NW of Pinos Altos, Hewitt 171 (GH); 
Guayanopa Canyon, Sierra Madre, Jones s.n. (POM, US); Santa Clara 
Mts., LeSueur 340 (ARIZ, CAS, GH, LL, TEX, UC); Mapula Mts., Pringle 
764 (LL, NY, RSA, UC, US). SONORA. La Mesa Colorado, Gentry 541, 
541m (ARIZ, DS); Sierra de Papas, Gentry 630 (Ds). 

Everly (1947) treated Perityle microcephala as a syn- 
onym of P. spilanthoides. I have found, however, that P. 
microcephala represents a valid species, while P. spilan- 
thoides is synonymous with the weedy variety of P. micro- 

Perityle microcephala belongs with the group of white- 
rayed species which includes P. turneri, P. lineariloba, and 
P. microglossa. Allof these taxa are quite similar in achene, 
pappus, and most other floral characters, but P. micro- 
cephala is clearly delimited from P. turneri and P. lineari- 
loba by its capitulescence of small, tightly clustered heads, 
grayish pubescence, and is easily separated from P. micro- 
glossa by its pubescence and perennial habit. Overall 
morphological similarity and distributional considerations 
(Fig. 3) suggest that P. microcephala is most closely re- 
lated to P. microglossa. Pringle 764 and Gentry 630, here 
ascribed to P. microcephala, approach P. microglossa in 
vegetative characteristics. 

9. Perityle microglossa Benth. Bot. Sulph. 119. 1844. TYPE: 
Nicaragua (?): Realejo, Nicaragua (?), 1837, R. B. Hinds 

9a. Perityle microglossa var. microglossa 

Perityle acmella Harv. & Gray, Mem. Amer. Acad. II. 4: 
77. 1849. TYPE: Mexico: “California”, Coulter 278 (Holo- 
type, GH!; isotypes, NY!). 

Galinsogeopsis spilanthoides Schultz-Bip. in Seem, Bot. 
Herald 307. 1856. TYPE: Mexico: SINALOA: Sierra Madre, 
1849, B. Seeman 1982 (Lectotype, K!). LECTOPARATYPE: 
same data, B. Seeman 1981 (K). 

Pericome spilanthoides Benth. & Hook. ex. Hemsl. Biol. 
Centr. Amer. Bot. 2: 215. 1881. 

Perityle microglossa var. effusa A. Gray, Syn. Fl. N. 

1974] Perityle — Powell 271 

Amer. 1: 332. 1884. TYPE: Arizona: Santa Catalina Mts., 
5 June 1882, C. G. Pringle (Holotype, GH!; fragment, us!; 
isotype NY!). 

Perityle effusa Rose, Contr. U.S. Nat. Herb. 1: 104. 1891. 
TYPE: Mexico: SONORA: Alamos, 26 Mar.-8 Apr. 1890, E. 
Palmer 350 (Lectotype, US!; isolectotypes GH !, NY !). LECTO- 
PARATYPE: same data, E. Palmer 377 (US!) ; isolectopara- 
types, (GH, NY !). 

Perityle spilanthoides (Schultz-Bip.) Rydb. N. Amer. FI. 
34: 17. 1914. 

Plants weedy, herbaceous annuals, 20-60 cm high, erect 
or decumbent, mostly branching above the base, stems few 
or many, upper portions, especially peduncles, copiously 
glandular-puberulent, rarely merely puberulent with few 
glandular hairs; leaves opposite or alternate, mostly oppo- 
site, puberulent, glandular-puberulent or glabrous, often 
turning purplish, (2.0) 3.0-7.0 (10.0) em long, (1.5) 2.0- 
5.0 (9.0) em wide, extremely variable in size and shape, 
cordate, ovate, broadly ovate-cordate to subreniform, or 
subdeltoid, the margins merely singly or doubly crenate- 
even or irregular, to strongly 3-lobed or cleft, pedately 
divided, or subhastate; petioles 0.5-3.0 (4.0) em long; capit- 
ulescence of 1-3 or many heads clustered on relatively 
short peduncles 0.6-3.5 (6.0) em long; heads radiate, 3.5- 
4.5 (6.0) mm high, 4.0-6.5 (7.0) mm wide, involucres cam- 
panulate; ray flowers 6-12, ligules white, 1.5-3.5 mm long, 
oblong; disc corollas yellow, 1.2-2.2 mm long, throats tubu- 
lar-funnelform; achenes 1.5-2.0 mm long, linear-oblong to 
linear-elliptic, rarely very narrow and short obovate, with 
thin to prominent callous margins, the margins ciliate, the 
faces glabrous or puberulent; pappus of a crown of squa- 
mellae and 2 unequal bristles, the longest 0.8-1.2 mm long; 
chromosome numbers, n = 34, 51. 

Rather widespread weed of northwestern Mexico, most 
common in Sonora, Sinaloa, and Baja California Sur, rang- 
ing south to Colima, seemingly introduced in northeastern 
Mexico and extreme south Texas. Flowering probably year 
around. (Fig. 3). 

272 Rhodora [Vol. 76 


MEXICO: Baja California Sur. San Jose del Cabo, Anthony 326 
(CAS, DS, GH, POM, UC, US) ; ca. 6 km NW of Mira Flores, Carter 2663 
(ns, UC, US) ; Arroyo de los Pozos, Moran 6886 (sp); La Paz, Palmer 
92 (CAS, GH, NY, US); Boca de la Sierra, Powell and Sikes 1672 
(SRSC, TEX); 1 mi N of Pescadero, Powell and Sikes 1676 (SRSC, 
TEX); vicinity of San Jose del Cabo, Wiggins 5689 (DS, NY, UC, US). 
Chihuahua. Tres Hermanos, SW of Batopilas, Hewitt 36 (GH). 
Colima. Colima, Oreutt 4551 (ps). Nayarit. Acaponeta, Jones s.n. 
(PoM, UC); San Blas, Maltby 22 (us); Maria Magdalena Islands, 
Maltby 175 (Ny, US); Tepic, Palmer s.n. (us). San Luis Potosi. 
near waterfall at El Salto, King 3877 (NY, TEX, UC, us); from San 
Luis Potosi to Tampico, Palmer 1093 (GH, NY, US). Sinaloa. Culiacan, 
Brandegee s.n. (UC); ca. 64 mi S of Culiacan, Breedlove 1545 (bs); 
Imala, Gentry 5455 (ARIZ, DS, NY, UC); San Blas, Jones, s.n. (CAS, NY, 
POM, UC); Lodiego, Palmer 1614 (GH, NY); 56 mi NW of Mazatlan, 
Powell and Sikes 1682 (sRsc, TEX); Rio del Fuerte, near El Fuerte, 
Rose, Standley and Russell 13587 (NY, US); vicinity of Villa Union, 
Rose, Standley and Russell 13932 (NY, US); ca. 5 mi E of Costa Rica, 
Sikes and Babcock 192 (SRSC, TEX); 70 mi S of Mazatlan, Sikes and 
Babcock 206 (SRSC, TEX). Sonora. 15 mi NE of Alamitos, Rio San 
Miguel, Abrams 13357 (ps); Hermosillo, Brandegee s.n. (DS, GH); 
12 mi W of Navojoa, Gentry 7950 (uc, US); Alamos, Palmer 673 
(us); Rio Mayo, Navojoa, Sikes and Babcock 177 (SRSC, TEX); 
2.4 mi N of Soyopa, Sikes and Babcock 157 (SRSC, TEX); 28 mi E 
of Navojoa, Sikes and Babcock 185 (SRSC, TEX). Zacatecas. near 
San Juan Capistrano, Rose 2427 (Ny, US). U.S.A.: TEXAS: Came- 
ron Co. Rio Hondo, Chandler 7007 (us); E of Rio Hondo, Clover 
1742 (ARIZ, TEX); Laguna Atacosa Refuge, Fleetwood 6069 (TEX). 
Willacy Co. Sauz Ranch, Johnston 53, 280 (TEX). 

The type locality originally given for Perityle m icroglossa, 
Realejo, Nicaragua, is probably in error. The southern-most 
eollection I have seen was from Colima, Mexico (Fig. 3), 
and it seems likely that the species would have been col- 
lected again in Nicaragua or in intermediate localities if it 
were there. The type locality probably was San Blas, Naya- 
rit, which also was visited during the Voyage of the Sulphur 
and where P. microglossa is known to occur. According to 
Rogers MeVaugh (personal communication), the localities 
of many other Sulphur collections were mixed up in similar 

The variable nature of Perityle microglossa is evident 

1974] Perityle — Powell 213 

from the synonymy and from Everly's (1947) confused 
treatment of the taxon. The current studies have led to the 
recognition of two biological entities within P. microglossa. 
One is var. microglossa (Fig. 3), a widespread, weedy poly- 
ploid (n = 34, 51), and the other is var. saxosa (Fig. 3), 
a more restricted "native" diploid (n — 17). Greenhouse 
tests utilizing numerous collections of var. microglossa have 
shown that it is self-compatible and that it reproduces 
prolifically by seed. Only a few such tests have been 
conducted with var. saxosa, but all have indicated self-in- 
compatibility for the diploid taxon. The morphological char- 
acters which distinguish the two varieties are not entirely 
consistent, and several morphological intermediates, for 
which chromosome numbers are not known, are found 
among existing collections. Variety microglossa typically 
exhibits shorter ligules, glandular-pubescent and shorter 
peduncles, tighter capitulescences, and a weedy habit, as 
compared to var. saxosa that usually displays longer ligules, 
rather smooth and longer peduncles, more lax capitules- 
cences, and a “native” habit. 

Morphologically, Perityle microglossa (particularly var. 
saxosa) is close to P. microcephala. Considering the evi- 
dence presented for these two species, it can be postulated 
that P. microglossa evolved from the montain-dwelling P. 
microcephala through derivation of annual habit and adap- 
tation to soil habitats. Variety microglossa must have 
evolved in response to polyploidization and development of 
reproductive self-compatibility which allowed for its 
“weedy” proliferation. 

Additional studies of Perityle microglossa var. saxosa, as 
well as P. microcephala, are most desirable. If var. saxosa 
proves to be a “native,” diploid, and self-incompatible pro- 
genitor of var. microglossa, then perhaps specific status 
should be accorded the latter taxon. 

9b. Perityle microglossa var. saxosa (Brandeg.) Powell, 
comb. nov. 

Perityle saxosa Brandeg. Zoe 5: 225. 1905. TYPE: Mexico: 
SINALOA: vicinity of Culiacan, Cerro Colorado, 5 Nov. 1904, 

274 Rhodora [Vol. 76 

Brandegee (Holotype, UC!; isotypes, GH!, POM!, US). 
Perityle urticifolia Rydb. N. Amer. Fl. 34: 15. 1914. 

TYPE: Mexico: SINALOA: Tepic, 5 Jan.-6 Feb. 1892, E. 

Palmer 1960 (Holotype, NY !; isotypes, GH !, UC!, Us!). 

Plants herbaceous annuals, possibly rarely weak peren- 
nials, 15-50 cm high, erect, upper stems and peduncles 
sparsely to densely puberulent, rarely moderately glandular- 
puberulent; leaves opposite or alternate, mostly opposite, 
glabrous to puberulent, rarely glandular-puberulent, 2-6 
(8) cm long, (1.0) 1.5-3.5 (5.0) em wide; capitulescence 
usually open, of 1-3 heads on relatively long peduncles, 2-7 
em long; ligules 3.5-4.5 mm long; dise corollas 1.7-2.0 mm 
long; achenes 1.0-1.6 mm long; pappus bristles 2, unequal, 
the longest 1.0-2.0 mm long; chromosome numbers n = 17, 

Mostly in foothills and lower elevations, both sides of the 
Sierra Madre Occidental, perhaps most common in Chihua- 
hua, Sonora, and Sinaloa, Flowering probably year around. 
(Fig. 3). 


MEXICO: Chihuahua. Temores, Flyr 60 (TEX); near Descanso, 
Flyr 77 (TEX); Palmer 238 (GH, NY, Us). Durango. Tamazula, 
Gentry 5234 (ARIZ, DS, GH, NY, UC). Nayarit, 4 mi E of Jalcocotan 
on road to Tepic, MeVaugh 12147 (Us). Sinaloa. 68 mi N of Ma- 
zatlan, Flyr 128 (TEX); 4 mi S of La Cruz junction, on MEX 15, 
Flyr 122 (TEX); Cerro Colorado, Gentry 5468 (ARIZ, DS, NY, UC, US); 
Canyon de Tarahumare, Sierra Surotato, Gentry 7315 (GH, NY, US); 
road from Las Flechas to La Rastra, Goldman 323 (GH, NY, US); 
30.4 mi E of Villa Union, Powell and Turner 1853 (SRSC, TEX). 
Sonora. just S of Jecori, Drouet, Richards, and Lockhart 3692 (ps, 
GH, NY); 18 mi NW of Campas, Drouet, Richards, and Lockhart 3704 
(Ds, GH); San Bernardo, Rio Mayo, Gentry 1255 (ARIZ, GH, UC); 
near Alamos, Gentry 4825 (ARIZ, DS, GH, NY); 12 mi SE of Colonia 
Oaxaca, Hastings and Turner 65-43 (ARIZ, DS, SD); Petaquilla Canyon, 
White 3320 (ARIZ, GH); La Vega Azul, SW of Cononia Morelos, White 
4852 (ARIZ, GH, NY, US); 35 mi NE of Cajeme, on road to Tesopaco, 
Wiggins 6408 (Ds, US). 

The Hastings and Turner and the White collections from 
NE Sonora are tentatively assigned to var. saxosa, but their 

1974] Perityle — Powell 275 

somewhat doubtful affinity with this taxon should be noted. 
The specimens have elongated achenes, approaching the 
fruit characteristics of Perityle ciliata and P. coronopofolia, 
and they are perennial. In vegatative features they are 
more like P. microglossa. Perhaps the unusual collections 
are from a relict population which is transitional between 
the shorter-achened (e.g., P. microglossa) and the longer- 
achened (e.g., P. ciliata) members of the white-rayed al- 
liance. Again, however, as is the case with most of the 
poorly collected Sierra Madre taxa, further study is re- 

10. Perityle canescens Everly, Contrib. Dudley Herb. 3: 393. 
1949. TvPE: Mexico: SINALOA: Capadero, Sierra Tacuicha- 
mona, 12 Feb. 1940, H. S. Gentry 5588 (Holotype, DS!; iso- 
types ARIZ!, GH !, NY !, UC], Us!). 

Plants low, suffruticose perennials, 6-15 cm high, densely 
leafed upper stems pilose; leaves opposite or alternate, 
pilose-hirsute to villous, 1.2-1.7 em long, 0.5-0.9 cm wide, 
3-pinnatifid with linear segments, ovate-cordate in outline, 
crisped; petioles 5-9 mm long; capitulescence essentially of 
solitary heads on short peduncles; heads radiate, 5-7 mm 
high, 6-8 mm wide, involueres broad campanulate; ray 
flowers ca. 10, ligules white, 2.5-3.5 mm long, oblong to 
broadly so; disc corollas yellow, becoming purple-tinged ram long, throats broad tubular; achenes 2.0-3.5 mm 
long, oblong-oblanceolate and nearly truncate on both ends, 
with thick callous margins, the margins merely puberulent, 
faces evenly puberulent; pappus of a prominent crown of 
united, erose squamellae, and 2 very unequal, rather stout, 
bristles, the longest 1.5-2.5 mm long; chromosome number, 

Known only from the type collection. 

Perityle canescens is related to P. coronopifolia from 
which it is distinguished by habit; dense indument; tend- 
ency for solitary heads; leaf morphology ; thickly calloused 
achenes with short-pubescent margins; strong, more or less 

276 Rhodora [Vol. 76 

united crown of pappus squamellae; stout pappus bristles: 
and distribution. 

Two other species, Perityle ciliata and P. hofmeisteria 
are grouped with the above taxa in the white-rayed alliance, 
largely on the basis of their similar floral characteristics 
(particularly achene and pappus). In habit and achene 
shape, P. canescens resembles P. lemmoni (sect. Laphamia: 
Powell, 1973), but the former taxon obviously belongs with 
sect. Perityle on the basis of its pappus crown. 

11. Perityle ciliata (L. H. Dewey) Rydb. N. Amer. Fl. 34: 
17. 1914. 

Laphamia ciliata L. H. Dewey, Bot. Gaz. 20: 425. 1895. 
TYPE: Arizona: rocks along Pine Creek near Pine, Ariz., 
26 Aug. 1891, D. T. MacDougal 676 (Holotype, US!; iso- 
type, US!). 

Plants suffruticose perennials, 15-30 cm high, many- 
stemmed, erect to pendulous, densely short-pubescent above; 
leaves opposite or alternate, densely gray-pubescent under- 
neath, pubescent to puberulent on upper surfaces, 0.8-2.0 
(3.0) em long, 0.6-1.3 (2.4) cm wide, deltoid-ovate to ovate- 
rhombic, the marigns serrate, shallow-lobed, or serrate- 
crenate; capitulescence of several heads clustered on short 
peduncles; heads radiate, 5-7 mm high, 5.5-7.0 mm wide, 
involucres campanulate; ray flowers ca. 6-10, ligules white, 
3-6 mm long, broadly oblong to oblong-elliptic; dise corollas 
yellow, often purple tinged, 2.0-2.5 (3.0) mm long, throats 
tubular to tubular-funnelform; achenes 2.0-2.8 mm long, 
linear-oblong, with prominent callous margins, the margins 
long-ciliate, the faces slightly angled and pubescent; pappus 
of a crown of squamellae and 2 (rarely 3) slender bristles, 
1.5-2.0 (2.5) mm long; chromosome number, n — 17. 

Restricted in distribution to the mountains of central 
Arizona; rock-dwelling. Flowering spring, summer, and 
fall. (Fig. 3). 

Arizona: APACHE C0.: Springerville, MeGinnies (ARIZ). COCONINO 
CO.: Long Valley, Coconino Natl. Forest, Coconino Rec. Crew 492 

1974] Perityle — Powell 277 

(us); Oak Creek, W of Troutdale, Goodding 198-47 (ARIZ, SMU); 
4 mi W of Strawberry, Sikes 99 (SRSC, TEX). GILA CO.: Barnhart 
Pass, Matzatzal Mts., Collom 102 (GH, NY, US); Fossil Creek Hill, 
Collom 594 (us); Parker Creek Canyon, Sierra Ancha Mts., Gould 
8632 (ARIZ, CAS, UC); Workman Creek Falls, Sierra Ancha, Johnson 
(ARIZ); Peterson Ranch, Sierra Ancha, Pase 1306 (ARIZ); 5.8 mi W 
of Strawberry, Sikes and Patterson 427 (SRSC, TEX). MOHAVE CO.: 
Hualapai Mts., Braem (ns); trail to Potato Patch, Hualapai Mts., 
Braem 875 (Ds). YAVAPAI CO.: near Senator Mine, along road from 
Prescott, Eastwood 16721 (CAS); between Prescott and Ash Fork, 
Eastwood 16770 (cas); Groom Creek, mts. near Prescott, Kearney and 
Peebles 9750 (ARIZ, uc); near Granite Dells, vicinity of Prescott, 
Kearney and Peebles 12780 (ARIZ, GH, NY, us); Prescott, Peebles, 
Harrison, and Kearney 2645 (ARIZ, US). 

This species is clearly related to P. coronopifolia from 
which it is conveniently delimited by leaf shape. 

12. Perityle coronopifolia A. Gray, Pl. Wright. 2: 82. 1853. 
TYPE: New Mexico: GRANT CO.: copper mines of Santa Rita 
del Cobre, Sept., 1851, C. Wright 1196 ( Holotype, GH!; 
isotypes, NY!, US!). 

Laphamia coronopifolia (A. Gray) Hemsl. Biol. Centr. 
Amer. Bot. 2: 210. 1881. 

Laphamia scopulorum M. E. Jones, Contr. West. Bot. 12: 
48. 1908. TYPE: Mexico: CHIHUAHUA: Colonia Juarez, 6000 
ft., 12 Sept. 1903, M. E. Jones (Holotype, POM!; fragment, 

Plants low, suffruticose perennials, 6-36 cm high, erect or 
pendulous, many-stemmed, very leafy, grayish-pubescent ; 
leaves opposite or alternate, puberulent to densely so, or 
nearly glabrous, 0.7-2.5 (3.5) cm long, 0.5-1.2 (2.0) cm 
wide, 3-palmate and lobed with slender-spathulate seg- 
ments; pedately divided, or delicately 2,3-pinnatifid with 
linear-filiform segments; petioles 2-8 (12) mm long; capi- 
tulescence of several heads clustered on short peduncles; 
heads radiate, 5.0-6.5 mm high, 5-6 mm wide, involucres 
campanulate; ray flowers ca. 8-12, ligules white, 3-7 mm 
long, broadly oblong, oblong-elliptic, to subspathulate; disc 
corolas yellow, often purple tinged, 2.0-2.8 mm long, 
throats tubular, tubular-funnelform, or tubular-campanu- 

278 Rhodora [Vol. 76 

late; achenes 1.8-2.5 mm long, linear-oblong to narrowly 
oblanceolate, with thin or prominent callous margins, the 
margins typically long-white ciliate, rarely short ciliate, the 
faces slightly angled and glabrous or pubescent; pappus of 
a crown of squamellae and 2 (rarely 3) slender bristles, 
1.5-2.5 mm long; chromosome number, n = 17. 

Rather widsepread and common in southern New Mexico- 
Arizona, and rare in northern Chihuahua, Mexico ; rock 
dwelling. Flowering spring, summer, and fall. (Fig. 3). 


MEXICO: Chihuahua. near Colonia Juarez, 6000 ft, Sierra Madres, 
Jones (POM, US). 

U.S.A.: Arizona: COCHISE co.: Miller Canyon, Hauchuca Mts., 
Barneby 5172 (CAS, NY); Ramsey Canyon, Huachuca Mts., Goodding 
758 (GH, NY, RSA). GRAHAM co.: Fry Canyon, Pinaleno Mts., Shreve 
1365 (ARIZ); Graham Mt., Thronber and Shreve 7783 (ARIZ). GREEN- 
LEE CO.: Mts. back of Clifton, Greene (NY); San Francisco Mts., 
Greene (POM); 0.5 mi N of Metcalf, Maguire, Richards and Moeller 
11822 (ARIZ, GH, NY, US). PIMA CO.: Spud Ranch, Rincon Mts., 
Blumer 3331 (ARIZ, DS, GH, UC) ; north slope Baboquivari Peak, Clark 
12565 (GH); San Pedro Vista, Santa Catalina Mts, Parker 8096 
(ARIZ, NY, RSA, US); below Goose-Head Rock, Mt. Lemmon, Sikes 95 
(SRSC). SANTA CRUZ co.: Flux Canyon, Patagonia Mts., Kearney 
and Peebles 10193 (ARIZ, US); Nogales to Ruby, Kearney and Peebles 
14961 (ARIZ, GH, NY). New Mexico: DOÑA ANA CO0.: W side, Organ 
Mts., Dunn 7274 (NMC, RSA); E peaks, Organ Mts., Powell 1393 
(SRSC); Dripping Springs Canyon, Organ Mts., Sikes 108 (SRSC). 
OTERO CO.: Dry Canyon, Rehn and Viereck (US). SOCORRO co.: 
Mogollon Creek, Mogollon Mts., Metcalf 323 (NMC, NY, UC, US); 
Gila Hot Springs, Mogollon Mts., Metcalf 828 (POM, UC, US). 

Perityle coronopifolia is closely related to P. ciliata, from 
which it is best distinguished by leaf Shape. The species 
also has affinity with P. canescens. Both P. coronopifolia 
and P. canescens have lobed leaves which are technically 
distinguishable, and they are further delimited by the long 
white-ciliate achene margins, clustered heads, pubescence, 
and distribution. 

The principal distribution of Perityle coronopifolia is in 
southern Arizona and New Mexico. The one known Mexican 
collection, originally described as Laphamia scopulorum, is 

1974] Perityle — Powell 279 

considered to be a morphological variant. The Jones speci- 
mens from near Colonia Juarez are both of poor quality 
but show delicate pinnatifid leaves, short-pubescent achene 
margins, and inconspicuous pappus squamellae. These two 
fruit characteristics indeed approach those of Laphamian 
taxa (Powell, 1973), but the strong indication of its rela- 
tionship with the Peritylean P. coronopifolia suggests its 
correct placement with the latter species. 

13. Perityle hofmeisteria Rydb. N. Amer. Fl. 34: 18. 1914. 
TyPE: Mexico: DURANGO: vicinity of Durango, Apr.-Nov., 
1896, E. Palmer 28 (Holotype, NY !; isotypes, GH !, US!). 

Plants suffrutescent to herbaceous perennials (possibly 
also annuals), 12-25 cm high, puberulent; leaves opposite 
or alternate, puberulent, 1.0-5.5 em long, 0.2-2.0 em wide, 
ovate to subspathulate and entire to shallow-lobed, or ovate- 
cordate and serrate to serrate-lobed ; petioles 0.5-2.0 (3.0) 
em long; capitulescence of 1-3 heads borne on relatively 
short peduncles, 1.5-2.8 cm long; heads radiate, 6-7 mm 
high, 4-6 mm wide, involucres turbinate-campanulate; ray 
flowers ca. 10-12, ligules white, 4-5 mm long, oblong; disc 
corollas yellow, 2.5-3.0 mm long, throats tubular-funnel- 
form; achenes 2.5-3.0 mm long, linear-oblong to narrowly 
obovate, with thin or thick callous margins, the margins 
prominently ciliate, the faces puberulent especially in the 
centers; pappus of a crown of squamellae and 2 subequal 
bristles, 2-3 mm long; chromosome number, n= 16 =+ 1. 

Seemingly endemic in Durango, Mexico; probably rock- 
dwelling. Flowering spring-fall. (Fig. 3). 

Mexico: DURANGO: 3 mi N of Rodeo, Flyr 332 (TEX). 

The Flyr 332 specimen, said by the collector to be *seem- 
ingly an annual," conforms to the types of Perityle hofmeis- 
teria in floral but not in vegetative features. The main dif- 
ferences are in habit appearance and leaf shape. The spe- 
cies is poorly understood at present, but its affinity is with 

280 Rhodora [Vol. 76 

P. ciliata and P. coronopifolia from which it is readily 
distinguished by leaf morphology and distribution (Fig. 3). 

14. Perityle jaliscana A. Gray in S. Wats. Proc. Amer. Acad. 
22: 431. 1887. TYPE: Mexico: JALISCO: Rio Blanco, ca. 10 
mi NW of Guadalajara, 17-23 Sept. 1886, E. Palmer 554 
(Holotype, GH ! ; isotypes, NY !, Us (2) !). 

Plants low, suffruticose perennials, 5-17 cm high, stems 
Spreading, densely puberulent; leaves opposite or alternate, 
densely puberulent to nearly glabrous, 2.0-3.5 em long, 0.8- 
1.8 (2.0) em wide, deltoid, deltoid-rhombic, deltoid-ovate, 
or subhastate to subhalberd, usually 2-4 lobed proximally, 
rarely serrate-lobed; petioles 0.7-1.8 (1.7) em long; capit- 
ulescence of 1-several heads clustered on short, slender 
peduncles; heads discoid, 5.0-6.5 mm high, 4-5 mm wide, 
involucres funnelform-campanulate; disc corollas cream- 
white and purple-tinged, 2.0-2.5 (3.0) mm long, throats 
tubular-campanulate; achenes 1.8-2.2 (2.8) mm long, ob- 
long or nearly so, slightly tapering to the base, truncate, 
with prominent callous margins, rarely with 3-callous mar- 
£ins, angled on both faces, the margins and faces short- 
pubescent; pappus of a vestigial erown of squamellae, and 
2 (3) slender bristles, 0.5-1.0 mm long, the bristles rarely 
flattened ; chromosome number, n = 17. 

Rock-dwelling endemic in Jalisco, Mexico. Flowering 
spring-fall. (Fig. 4). 


Mexico: JALISCO: ca. 40 km N of Guadalajara, road to San Cristo- 
bal de la Barranca, MeVaugh 22114 (NY, sD); Sierra San Esteban. 
near Guadalajara, Pringle 2352 (GH, UC, US) ; Sierra de San Esteban, 
Pringle 156834 ( ARIZ, CAS, LL, SMU, US); Rose and Painter 7476 (NY, 
US); 5 mi SW, by road, from San Cristobal de la Barranca, Sikes 
and Babcock 383 (SRSC, TEX). 

Although relatively few collections of Perityle jaliscana 
are known, it is the best-collected of four closely related 
endemic species of Jalisco, Mexico. The others, P. rosei, 
P. trichodonta, and the recently described P. feddemae, are 
known only from type collections. 

1974] Perityle — Powell 281 

jaliscana (open circles); P. rosei 
(open square); P. feddemae (closed 

Fig. 4. Distribution of P. 
(closed square); P. trichodonta 


282 Rhodora [Vol. 76 

Perityle jaliscana and P. feddemae are the most distinct 
of the four species. I have followed Everly (1947) in recog- 
nizing P. rosei and P. trichodonta, even though future col- 
lections might indicate that they should be merged as a 
single species. Furthermore, both P. rosei and P. tricho- 
donta might simply be ecological variants of P. jaliscana. 
I recognize the four Species at this time mainly to preserve 
taxonomic consistency. In all three sections of the genus 
Perityle there are groups of closely related species that are 
no more distinctive than are the Jalisco endemics. In 
Jalisco, P. feddemae, with its dissected leaves, seems to be 
quite distinct, and it was recently discovered in a locality 
not far from the other species (Fig. 4). Strict endemism 
is common among the rock-dwelling taxa of Perityle, and 
lacking evidence to the contrary, I think it possible that the 
taxa in question are endemic species. 

Perityle jaliscana is most closely related to P. rosei. Most 
of its distinctive features, such as discoid heads, smaller, 
oblong achenes, shorter pappus bristles and squamellae, 
subhastate leaves, and short, dense pubescence, appear as 
if they could have been derived from P. rosei. 

15. Perityle rosei Greenm. Proc, Amer. Acad. 40: 45. 1905. 
TYPE: Mexico: JALISCO: in the Sierra Madre W of Bolanos, 
15-17 Sept. 1897, J. N. Rose 2947 (Holotype, Us!; isotypes, 
GH !, NY!). 

Plants suffruticose perennials, 10-20 em high, stems erect 
or spreading, densely short-pilose; leaves opposite or alter- 
nate, short-pilose to puberulent, 1.8-3.0 em long, 1.0-1.5 
(2.0) em wide, deltoid to deltoid-ovate or deltoid-rhombic, 
rarely subhastate, the margins entire or nearly so; petioles 
5-9 mm long; capitulescence of 1.-several heads on short 
peduncles; heads radiate, ca. 5 mm high, 4-5 mm wide, 
involucres campanulate; ray flowers ca. 13, ligules white, 
2.0-2.5 mm long; dise corollas cream-white and purple- 
tinged, 2.0-2.3 mm long, throats tubular-campanulate ; 
achenes 2.0-2.5 mm long, narrowly obdeltoid, with promi- 
nent callous margins, slightly angled on both faces, the 

1974] Perityle — Powell 283 

margins and faces puberulent to subglabrous; pappus of a 
short crown of squamellae, more or less united, and 2 (3-4) 
slender bristles, 1.5-2.0 mm long, the 3-4 bristles, if present, 
reduced in length; chromosome number, unknown. 

Known only from the type collection. (Fig. 4). 

Perityle rosei is best delimited from P. jaliscana by its 
radiate heads, nearly obconical achenes, longer crown of 
pappus squamellae and longer bristles, nearly deltoid, sub- 
entire leaves, and longer, less dense pubescence. 

16. Perityle trichodonta Blake, Proc. Biol. Soc. Wash. 37: 
60. 1924. TYPE: Mexico: JALISCO: Sierra Madre W of Bo- 
lanos, 15-17 Sept. 1897, J. N. Rose 2978 (Holotype, us!; 
isotype, GH !). 

Plants low, suffruticose perennials, ca. 10 cm high, many- 
stemmed and spreading, densely gray-pubescent above; 
leaves mostly opposite, densely hirtellous-pilose and gray- 
ish, 7-9 mm long, 2.5-5.0 mm wide, ovate to ovate-rhombic 
and subhastate; petioles 2-4 mm long; capitulescence of 
1-several heads clustered on short peduncles; heads radiate, 
ca. 4.5 mm high, ca. 4 mm wide, involucres funnelform- 
campanulate; ray flowers ca. 8, ligules white, 1.0-1.5 mm 
long; disc corollas cream-white and purple-tinged, 2.0-2.5 
mm long, throats tubular-campanulate; achenes 1.5-2.2 mm 
long, narrowly obdeltoid, with prominent callous margins, 
slightly angled on both faces, the margins and faces puberu- 
lent to subglabrous; pappus of a short crown of squamellae, 
more or less united, and 2 slender bristles, 1.5-2.2 mm long, 
chromosome number, unknown. 

Known only from the type collection. (Fig. 4). 

Perityle trichodonta looks as though it could be a small- 
leafed, more densely pubescent, shorter liguled form of P. 
rosei, but it does differ in the characters listed. Future col- 
lections might reveal that P. trichodonta should be merged 
with P. rosei. 

17. Perityle feddemae McVaugh, Contr. Univ. Mich. Herb. 

284 Rhodora [Vol. 76 

9: 431-433. 1972. TvPE: Mexico: JALISCO: Rancho Viejo, 
ca. 10 km ENE of Huejuquilla el Alto, rock-crevices, open 
oak-woodland, elev. 1900 m, 31: Oct. 1963, Feddema 2278 
(Holotype, MICH ; isotype, SRSC!). 

Plants suffruticose perennials, 10-25 cm high, stems 
spreading, puberulent; leaves mostly opposite, puberulent, 
glandular-dotted, 1.7-3.5 em long including petioles, dis- 
sected-pinnatifid with linear-spatulate segments; capitu- 
lescence of 1-3 heads on short, slender peduncles; heads 
discoid, 5-6 mm high, 3-5 mm wide, involucres funnelform- 
campanulate; disc corollas pale yellow, 2.3-2.8 mm long, 
throats tubular-campanulate; achenes 2-3 mm long, oblong- 
oblanceolate, with prominent callous margins, angled on 
both surfaces, the margins and faces short-pubescent; pap- 
pus of a short crown of squamellae and 2 slender, unequal 
bristles, ca. 1.0 mm long; chromosome number, unknown. 

Known only from the type collection. (Fig. 4). 

This taxon possibly is most closely related to Perityle 
jaliscana that also has discoid heads, but its dissected leaves 
mark it as the most distinctive of the Jalisco endemics. 

18. Perityle parryi Gray, Pl. Wright. 2: 106. 1853. TYPE: 
Texas: Canyon of the Rio Grande, Aug. 1852, C. C. Parry 
521 (Isotype, NY!). 

Laphamia parryi (Gray) Benth. & Hook. ex Hemsl. Biol. 
Centr. Amer. Bot. 2: 210. 1881. 

Leptopharyna trisecta Rydb. N. Amer. Fl. 34: 23. 1914. 
TYPE: Mexico: CHIHUAHUA: dry cliffs, rocky hills near 
Chihuahua, 20 Mar. 1885, C. G. Pringle 183 (Holotype, NY l; 
isotypes, DS!, GH !, UC!, US!). 

Plants low, woody-based perennials in crevices of rocks 
or tall, spreading, herbaceous to suffruticose perennials in 
soil, 10-75 em high; leaves opposite or alternate, glandular- 
pubescent, 2-6 (9) cm long, 0.8-4.0 (6.0) cm wide, cordate 
to subreniform in outline, margins irregularly lobed, den- 
tate, or laciniate, but typically shallow to deeply 3-lobed, 
and in some specimens even subcruciform; petioles 1.2- 

1974] Perityle — Powell 285 

Fig. 5. Distribution of P. parryi (open triangles); P. vaseyi 
(closed triangles); P. aglossa (open squares); P. cordifolia (closed 
circles); P. leptoglossa (closed squares); P. lloydii (open star); 
P. lobata (closed stars) ; P. palmeri (open circles). 

286 Rhodora [Vol. 76 

4.0 em long; capitulescence of 1-3 heads borne on pe- 
duncles 1-7 cm long; heads radiate (rarely discoid), 0.8-1.0 
em high, 0.7-1.4 em wide, involucres hemispherical to campa- 
nulate; bracts numerous, lanceolate to linear, rarely ob- 
lanceolate, acute to attenuate at the apex; ray flowers ca. 
12-16 (rarely 1-6), ligules yellow, oblong, showy; disc flow- 
ers numerous, corollas yellow, (3.2) 4-6 mm long, throats 
tubular; achenes (2) 3-4 mm long, linear-elliptic to ob- 
conical-elliptic, with thin to prominent callous margins, the 
margins short-pubescent to dense-pilose, faces pubescent ; 
pappus of a crown of squamellae, and 1 (rarely 0 or 2) 
barbellate bristle, (1.0-2.5) 3-6 mm long, typically tapering 
to a point, the barbs typically antrorse throughout, rarely 
becoming lateral or retrorse toward the tip; chromosome 
number, n = 17. 

Western Big Bend area of Texas and adjacent Mexico; 
occurring in rocks and soils. Flowering essentially year 
around. (Fig. 5). 


MEXICO: Chihuahua. 11 mi NE of Camargo, Johnston 7920 
(GH); road to San Carlos Mines, Johnston and Muller 39 (GH); 
12 mi S of Ojinaga, Johnston and Muller 1443 (GH); near new lake 
on Rio Conchos, Powell, Turner, and Magill 2030 (SRSC, TEX); rocky 
hills near Chihuahua, Pringle 183 (Ds, GH, NY, UC, US); 1 mi E of 
km 100, Ojinaga-Camargo road, Sikes and Patterson 397 (SRSC, TEX) ; 
20 km N of Chihuahua, Stewart and Johnston 2119 (GH). U.S.A.: 
Texas: Brewster Co. Lost Mine Peak, Chisos Mts, Correll and 
Johnston 24540 (LL); Green Valley Ranch, Correll and Wasshausen 
27825 (LL); Agua Fria Spring, Cory 18638 (GH); Santa Helena 
Canyon, Cory 26475 (us); N side Whirlwind Spring area, 02 Ranch, 
Fletcher 1143 (srsc); Packsaddle Mt., McAfee 261 (SRSC); Chisos 
Mts., Muller 8247 (GH, NY, US, TEX); lower Window Trail, Big Bend 
Park, Powell and Sikes 1528 (sRSC, TEX); Basin, Chisos Mts., Sikes 
32 (SRSC, TEX); Boot Springs Trail, Sikes 116 (SRSC, TEX); Nine- 
Point Mesa, Warnock and Johnston 17670 (srsC); Rosillos Mts., 
Warnock 20603 (sRsc). Presidio Co. S edge of Presidio, Correll and 
Johnston 21867 (LL); 10 mi SE of Redford, Correll and Johnston 
21898 (LL); Casa Piedra road, 20 mi S of Marfa, Cox s.n. (SRSC) ; 
Capote Creek, Havard s.n. (US); near Cottonwood Springs, Sierra 
Tierra Vieja, Hinckley 1850 (GH, NY, US); ca. 9 mi N of Lajitas, 
Powell 1243 (sRSC) ; 5 mi S of Redford, Powell, Sikes, and Watson 

1974] Perityle — Powell 287 

1396 (sRsc); 12 mi S of Redford, Powell, Sikes, and Watson 1398 
(SRSC); 3 mi E of Ruidosa, Powell and Brey 1513 (sRsc); Capote 
Creek, ca. 1 mi below Capote Falls, Powell 1518 (sRsC). 

The three yellow-flowered species of west Texas and ad- 
jacent Mexico (Fig. 5), Perityle parryi, P. vaseyi, and P. 
aglossa, are morphologically similar. Perityle parryi and 
P. vaseyi are particularly close and somewhat confusing 
taxonomically, primarily because of intermediate leaf forms 
and uncertain distributional integrity. 

Perityle parryi occurs as a rock-dwelling subshrub and as 
a soil-dwelling perennial, typically with leaves that are 
merely 3-lobed. Perityle vaseyi is a soil-dwelling, taprooted 
perennial, with deeply trisected leaves. However, some 
Mexican populations of P. parryi (— L. trisecta) have 
deeply trisected leaves, as do some soil-dwelling Texas 
forms, and it is not always possible to distinguish the taxa 
by leaf morphology. Perityle parryi is further delimited by 
long-tapering pappus bristles (one per achene) which are 
antrorsely barbellate, bracts usually lanceolate or linear and 
somewhat attenuate, and disc corollas with usually narrowly 
tubular throats, as opposed to P. vaseyi with shorter pappus 
bristles that are retrorsely barbellate at the tips, bracts 
usually oblanceolate and acute, and more broadly tubular 
dise corollas. 

It is suspected that the soil-dwelling forms of Perityle 
parryi and P. vaseyi may occasionally hybridize. Any hy- 
bridization, however, would be difficult to detect morpho- 
logically. It is not known whether the leaf variability of 
P. parryi is intrinsic or whether it might be the result of 
hybridization with P. vaseyi. In spite of marginal sympatry 
(Fig. 5), mixed populations of the two species have not 
been found, and no clear-cut hybrids have been detected. 
The subject is under experimental study. 

The populations of Perityle parryi near Ruidosa, Texas, 
in the Sierra Vieja and one collection 12 mi S of Ojinaga 
(Johnston and Muller 1443) are notably anomalous. Their 
pappus bristles are short and have retrorsely barbed tips, 
resembling those of P. vaseyi. In addition, the Sierra Vieja 

288 Rhodora [Vol. 76 

populations may be nearly rayless or entirely discoid. In 
other morphological characters the anomalous populations 
conform with P. parryi. The discoid forms should not be 
confused with P. aglossa, which is always rayless. 

19. Perityle vaseyi Coult. Contr. U. S, Nat. Herb. 1: 42. 
1890. TYPE: Texas: BREWSTER CO. Chisos Mts., 1889, G. C. 
Nealley (245?), (Holotype, Us!). 

Plants perennial with fleshy taproots, (10) 15-75 cm 
high, herbaceous to suffruticose, stems erect or spreading; 
leaves opposite or alternate, glandular-pubescent, 2.5-6.5 
em long, 2-5 em wide, typically palmately divided into 3 
lobes or subcruciform, with the lobes also dissected, cleft, 
or parted, rarely the leaves are strongly 3-lobed, cleft, or 
parted; petioles 1.4-3.5 cm long; capitulescence of 1-3 heads 
borne on peduncles 1-8 cm long; heads radiate, 0.8-1.0 cm 
high, 1.0-1.3 em wide, involucres hemispherical to campanu- 
late; bracts numerous, oblanceolate to linear-lanceolate, 
typically acute, less often alternate at the apex; ray flowers 
ca. 14-16, ligules yellow, oblong, showy; disc flowers numer- 
ous, corollas yellow, 3.2-5 (6) mm long, throats tubular to 
broadly so; achenes (2.0) 3.0-4.0 mm long, linear-elliptic 
to obconical-elliptic, with thin to prominent callous margins, 
the margins short-pubescent to dense pilose, faces pubes- 
cent; pappus of a crown of squamellae, and 1 (rarely 0 or 
2) barbellate bristle, 1.5-4.0 mm long, typically truncate, 
the barbs retrorse or lateral toward the tip; chromosome 
number, n — 17. 

Desert soils, especially gypsiferous clay, southwestern 
Big Bend area of Texas and adjacent Mexico. Flowering 
essentially year around. (Fig. 5). 


MEXICO: Chihuahua. 6.5 mi S of Ojinaga, Powell, Turner, and 
Magill 2002 (sRsC, TEX); 5 mi S of Ojinaga, Powell, Turner, and 
Magill 2079 (sRsc, TEX). Coahuila. Picachos Colorados, Johnston 
and Muller 141 (GH). U.S.A.: Texas: BREWSTER CO. petrified forest, 
Cory 2706 (GH); Hot Springs, Cory 6477 (POM); Glenn Springs, 
Cory 44014 (TEX); Terlingua Creek on 02 Ranch, Fletcher 917 

1974] Perityle — Powell 289 

(SRSC); 2-8 mi E of Terlingua, Flyr 16 (SMU, SRSC, TEX); 8 mi E 
of Lajitas, Flyr 19 (SMU, TEX); 1 mi SE of Terlingua, Flyr 180 
(SMU, TEX) ; 2 mi SE of Hen Egg Mt., Johnston and Warnock 3656 
(SRSC); top third of Packsaddle Mt., McAfee 184 (snsc); 70 mi S 
of Alpine, Powell and Sikes 1530 (sRSC, TEX); 18 mi SE of Castolon, 
Powell, Averett, and Watson 1539 (sRsC, TEX) ; 0.4 mi N of Castolon, 
Powell, Averett, and Watson 1544 (SRSC, TEX); near Big Bend Tun- 
nel, Sikes 39 (SRSC, TEX) ; near hwy. 118 entrance to Big Bend Park, 
Sikes 40 (SRSC, TEX); 7 mi E of Castolon, Sikes and Averett 330 
(SRSC, TEX); road to Fisk Canyon, Warnock 1148 (sRsc); Avary 
Canyon, Big Bend Park, Warnock 9139 (sRsC); Big Bend Park, 
Warnock and Wallmo 12211 (sRSC); San Vicente, near the Rio 
Grande, Young s.n. (TEX). 

As discused previously under Phylogenetic Considera- 
tions, an ancestral connection is hypothesized for Perityle 
vaseyi and its allies with the P. crassifolia group of Baja 
California. The primary evidence for this possible relation- 
ship are the similar habits and pappus bristles (with re- 
trorsely barbed tips) of P. vaseyi and P. crassifolia and the 
chromatographic profiles with yellow compounds, common 
to both groups. 

Morphologically, Perityle vaseyi and its relatives are 
closer to the P. cordifolia group of western Mexico. The 
distribution of the Texas species (Fig. 5) could have been 
achieved by migration from a Sierra Madre Occidental 
origin. The headwaters of the Rio Conchos are in the Sierra 
Madre near Creel, just across the divide from P. cordifolia 
country. The distribution of P. parryi near the Rio Conchos 
in eastern Chihuahua and the location of P. vaseyi and P. 
aglossa near the Rio Grande, suggest the possibility of 
eastern migration along the river systems. 

20. Perityle aglossa Gray, Pl. Wright 2: 107. 1853. TYPE: 
Texas: Rio Grande below Presidio del Norte, Aug. 1852, 
C. C. Parry (Lectotype, GH !). 

Laphamia aglossa (Gray) Benth. & Hook. ex Hemsl. Biol. 
Centr. Amer. Bot. 2: 210. 1881. 

Leptopharynx aglossa (Gray) Rydb. N. Amer. Fl. 34: 
24. 1914. 

Plants woody-stemmed perennials in rock crevices, 15-45 

290 Rhodora [Vol. 76 

cm high; leaves opposite or alternate, glandular-pubescent, 
2.4 em long, 0.6-2.3 cm wide, ovate, subeordate or subdel- 
toid, irregularly lobed, laciniate, or serrate-dentate, often 
shallow-3-lobed; petioles 0.8-2.3 em long; capitulescence of 
1-3 heads borne on peduncles 1.0-2.5 em long; heads discoid, 
8-11 mm high, 4-8 mm wide, involucres campanulate; bracts 
linear to linear-lanceolate, attenuated at the apex; disc 
flowers numerous, corollas yellow, typically pink- or purple- 
tinged, 4.2-6.0 mm long, throats tubular; achenes (2.0) mm long, oblong-obconical, with prominent callous 
margins, the margins short-pubescent to dense-pilose, faces 
pubescent; pappus of a crown of squamellae and 1 (rarely 
2) barbellate bristle, 3.7-5.5 mm long, tapering to a point, 
the barbs antrorse throughout; chromosome number, 
n = 17. 

Restricted in southern Brewster Co. and Terrell Co., 
Texas, along and near the Rio Grande; crevices of lime- 
stone. Spring-fall. (Fig. 5). 


Texas: BREWSTER CO.: mouth of Regan Canyon on Rio Grande, 
Hinckley and Warnock 3737 (US, SRSC); near Boquillas, Marsh. 306 
(GH); near Big Bend Tunnel, Sikes 38 (sRsC); 4 mi W of Hot 
Springs, Big Bend Park, Warnock and Turner 8291 (sRsC); San 
Vicente Canyon, Big Bend Park, Warnock and Parks 864 (LL, SMU, 
SRSC); limestone hills, Terlingua area, Warnock 13980 (SRSC) ; 
Heath Canyon, Black Gap Refuge, Warnock 20615 (sRSC); mouth 
of Regan Canyon, Warnock and Hinckley 461050 (SMU, SRSC, TEX). 
TERRELL CO.: mouth of San Francisco Canyon, Surratt 133 (SRSC) ; 
mouth of San Francisco Canyon, Warnock and Surratt 9817 (LL, 
SMU, SRSC) ; 9 mi E of Sanderson, Warnock 13155 (sRsC) ; along Rio 
Grande, between Regan Canyon and Sanderson Canyon, Warnock 
15854 (sRsC). 

Although only one type specimen is cited for Perityle 
aglossa (Parry, Aug. 1852), I have seen another specimen 
which probably came from Parry's original collection. At 
NY a sheet labeled *P. aglossa, Arroyo San Carlos, April 
1852," displays specimens of both P. aglossa and P. parryi. 
Judging from present-day distributions of the species, I 
believe that the specimens were probably mixed after 

1974] Perityle — Powell 291 

Parry's collections were made. Furthermore, the P. aglossa 
specimen on the NY sheet probably was from the original 
colleetion and perhaps should be recognized as an isolecto- 
type. The P. parryi specimen is discoid and probably came 
from the vicinity of Ruidosa above Presidio. 

Perityle aglossa is a limestone-dwelling subshrub with 
good distributional integrity. In addition to habit and dis- 
coid heads, the species is distinguished from P. parryi, its 
closest relative, by subtle characters of the leaves, involu- 
eral bracts, and disc corollas. 

21. Perityle cordifolia (Rydb.) Blake, Contr. U. S. Nat. 
Herb. 23: 1602. 1962. 

Leptopharynx cordifolia Rydb. N. Amer. Fl. 34: 22. 
1914. TYPE: Mexico: SINALOA: vicinity of Topolobampo, 
23 Mar. 1910, J. N. Rose, P. C. Standley, and P. G. Russell 
13294 (Holotype, NY!?; fragment of type, UC!; isotype, 

Plants suffrutescent perennials, rather low and spreading 
or semi-pendulent, 10-20 cm high, stems strong but herba- 
ceous except at the bases, pilose; leaves mostly alternate, 
bright green, pilose, 3.5-11.0 cm long, 1.5-9.0 cm wide, cor- 
date to broadly so, veins prominent, the margins doubly 
serrate-dentate, rarely crenate; petioles 1.5-4.5 mm long; 
capitulescence of solitary heads borne on rather stout pe- 
duncles, 2.5-6.0 em long; heads radiate, rarely cernuous, 
10-12 mm high, 9-15 mm wide, involucres hemispherical] to 
campanulate, typically truncate at the base; bracts 10-13 
mm long, linear and attenuate; ray flowers numerous, lig- 
ules yellow, 9-15 mm long, oblorg-elliptic; disc flowers 
numerous, corollas yellow, 5-7 mm long, throats tubular ; 
achenes 2.5-3.0 mm long, narrowly oblong, with thin or 
prominent callous margins, the margins typically densely 
ciliate, rarely puberulent, faces glabrous or puberulent in 
the center; pappus of a crown of squamellae, ca. 0.5 mm or 
less, and a single bristle, 4-7 mm long, the bristle tapering 
from base, a fine tip and densely barbellate; chromosome 
number, n = 17. 

292 Rhodora [Vol. 76 

A petrophilic, cliff-dwelling species, possibly also rooted 
in soils of shaded bluffs, Sonora and Sinaloa, Mexico. Flow- 
ering probably year around. (Fig. 5). 


Mexico: SINALOA: hill near Topolobampo, Flyr 63 (TEX); N of 
Topolobampo, Flyr 76 (TEX); road to Los Animas beach from Los 
Mochis, Flyr 105 (TEX); 23 mi N of Culiacan, Flyr 111B (TEX); 
Cerro Tecomate, W of Pericos, Gentry 5743 (ARIZ, DS, GH, NY, UC); 
ca. 1 mi E of Topolobampo, Sikes and Babcock 188 (SRSC, TEX); 
bluffs W of Topolobampo, Sikes and Babcock 190 (SRSC, TEX). 
SONORA: 32 mi N of Navojoa, Flyr 103 (TEx); Canyon Sapopa, 
Rio Mayo, Gentry 1282 (ARIZ, GH, NY, UC); Aquibiquichi, near Bata- 
cosa, Gentry 3003 (ARIZ, CAS, GH, UC, US); Cerro de Bayajori, 12 mi 
W of Navojoa, Gentry 7949 (uc, US); 23 km E of Navojoa, Moran 
4032 (DS, sD, UC); 1-2 mi N of San Carlos Resort, Powell and Sikes 
1689 (SRSC, TEX); 10 mi N of Guaymas, Randle and Werner 1 (ARIZ). 

Perityle cordifolia belongs with a group of four other 
morphologically similar species; P. palmeri, P. leptoglossa, 
and P. lloydii also of northwestern (mainland) Mexico, and 
P. lobata of Baja California. In spite of their similarities, 
the taxa are rather clearly delimited by vegetative features, 
excepting P. lloydii, which has characters of the other three 
mainland species. I have grown all of the taxa except P. 
lloydii under identical conditions, and each did maintain its 
characteristic morphology. 

Everly (1947) treated Perityle cordifolia as synonymous 
with P. palmeri. I recognize these taxa as related but cer- 
tainly distinct species. In fact, the soil-dwelling P. palmeri 
could have evolved from the rock-dwelling P. cordifolia, but 
on morphological grounds, P. palmeri shares even closer 
relationship with P. lobata. Perityle cordifolia is readily 
delimited from P. palmeri by its woody habit; bright green, 
pilose, cordate leaves with prominent veins and doubly ser- 
rate margins; long ligules; long attenuate involucral bracts; 
long pappus bristles; and long style branches. 

22. Perityle leptoglossa Harv. & Gray in A. Gray, Mem. 
Amer. Acad. II. 4: 77. 1849. TYPE: Mexico: SONORA: “vi- 
cinity of Hermosillo,” 1829-1830, Coulter 277 (Holotype, 
GH!; fragment, Us!). 

1974] Perityle — Powell 293 

Leptopharynx leptoglossa (Harv. & Gray) Rydb. N. 
Amer. Fl]. 34: 22. 1914. 

Plants shrubby, spreading or erect, 10-60 cm high, stems 
densely grayish-puberulent; leaves mostly alternate, densely 
puberulent, gray-green, 3.0-5.5 em long, 1.3-3.5 cm wide, 
cordate to cordate-ovate, the margins shallow-lobed and 
crenate to serrate, or merely crenate-serrate; petioles 1.3- 
2.5 cm long; capitulescence of solitary heads borne on rela- 
tively stout peduncles, 2-5 em long; heads radiate, rarely 
cernuous, 8-10 mm high, 7-12 mm wide, involucres cam- 
panulate-cylindroidal and truncate at the base; bracts 5-8 
mm long, linear to linear-lanceolate and attenuate to sub- 
attenuate; ray flowers numerous, ligules yellow, 6-10 mm 
long, oblong to oblong-elliptic; disc flowers numerous, corol- 
las yellow, 4-5 mm long, throats tubular; achenes (2.2) 
2.5-3.0 mm long, narrowly oblong, with thin or prominent 
callous margins, the margins long- or short-ciliate, the faces 
glabrous or minutely pubescent at the center; pappus of a 
crown of squamellae less than 0.5 mm long, and a single 
bristle, 2.5-4.5 mm long, the bristle tapering from base to 
apex and densely barbellate; chromosome number, n = 17. 

Strictly rock-dwelling, in the vicinity of Hermosillo, So- 
nora, Mexico. Flowering probably year around. (Fig. 5). 


Mexico: SONORA: 3 mi S of Magdalena, Aleramo 13216 (ps); 
rocky sides of El Cerro de la Campana, Drouet, Richards and Alva- 
rado 3387 (DS, GH, NY); 6 km N of Hermosillo, Drouet and Richards 
3757 (ps); San Miguel de Horcasitas, Eisen (UC); 41 mi N of 
Hermosillo, Flyr 96 (TEX) ; 26 mi S of Hermosillo, Gentry 45'12 (UC) ; 
ca. 28 mi N of Hermosillo, Powell and Averett 1520 (SRSC, TEX) ; 
ca. 28 mi N of Hermosillo, Powell and Sikes 1683 (SRSC, TEX); near 
Hermosillo, Ripley 14309 (CAS) ; vicinity of Hermosillo, Rose, Standley, 
and Russell 12346 (NY, US); near Magdalena, Shreve (NY); near 
Estacion Torres, Shreve 6070 (ARIZ); 8 mi W of Hermosillo, Wiggins 
and Rollins 96 (ns, GH, NY); ca. 1 mi N of Torres, Wiggins 6266 
(DS, GH, POM, US); 12 mi from Ures on road to Babiacari, Wiggins 
7359 (DS, TEX, US). 

The most distinctive features of Perityle leptoglossa are: 

294 Rhodora [Vol. 76 

shrubby habit; leaf margins shallow-lobed or merely cre- 
nate-serrate; densely puberulent upper stems and leaves; 
small heads and short ligules; short bracts; achene margins 
typically short-ciliate; pappus bristles typically slender at 
bases. At one locality ca. 28 mi N of Hermosillo, I observed 
a few plants that were 214 ft. high, the largest woody plants 
in the genus. The species is related to P. palmeri and P. 

23. Perityle lloydii Rob. & Fern. Proc. Amer. Acad. 30: 118. 
1894. TvPE: Mexico: SONORA. Badehuachi, 2 Dec. 1890, 
C. E. Lloyd 400 (Holotype, GH ! ; isotypes, NY !, UC!, US(3)!). 

Leptopharynz lloydii (Rob. & Fern.) Rydb. N. Amer. Fl. 
34: 22. 1914. 

Plants semishrubby, erect, 10-40 cm high, stems woody 
to herbaceous, pilose; leaves mostly opposite, pilose, 2.5- 
4.5 em long, 1.5-2.5 em wide, cordate to cordate-ovate, the 
margins doubly serrate; petioles 1.5-3.0 em long; capitu- 
lescence of solitary heads borne on peduncles 2.0-3.5 cm 
long; heads radiate, rarely cernuous, 8-9 mm high, 6-7 mm 
wide, involucres campanulate-cylindroidal and truncate at 
the base; bracts 5-7 mm long, linear and attenuate; ray 
flowers numerous, ligules yellow, 5-8 mm long, oblong to 
oblong-elliptic; disc flowers numerous, corollas yellow, ca. 
4 mm long, throats tubular; achenes ca. 3 mm long, nar- 
rowly oblong, the callous margins short-pubescent; pappus 
of a minute crown of squamellae (ca. 0.2 mm long), and a 
single bristle ca. 3 mm long, the bristle slightly tapering 
from base to apex and densely barbellate; chromosome 
number unknown. 

Known only from the type collection. (Fig. 5). 

Perityle lloydii is poorly delimited from the other main- 
land taxa of the P. cordifolia species-group. In leaf shape, 
pubescence, and achene morphology (short-pubescent mar- 
gins and reduced pappus squamellae) P. lloydii resembles 
P. cordifolia. In shrubby habit, small heads, short ligules, 
and other floral characters, the taxon approaches P. lepto- 

1974] Perityle — Powell 295 

glossa. Its pubescence is like that of P. palmeri. Overall, 
P. lloydii possibly is closer to P. leptoglossa, and further 
collections might well provide evidence for merging the 

24. Perityle lobata (Rydb.) I. M, Johnston, Proc. Calif. 
Acad. IV. 12: 1205. 1924. 

Leptopharynz lobata Rydb. N. Amer. Fl 34: 23. 1914. 
TYPE: Mexico: BAJA CALIFORNIA: Commondu, 19 Feb. 1889, 
T. S. Brandegee (Holotype, GH!; isotype, uc!). 

Plants fleshy-rooted perennials, possibly also annuals, 
low and spreading, semierect, or semipendulent, 10-40 cm 
high, stems weak, herbaceous, and rather brittle, usually 
purplish, glabrous to sparse-pilose ; leaves mostly alternate, 
bright green, turning purplish, glabrous to sparse-pilose, 
3-8 (12) cm long, 1.8-6.0 (8) cm wide, cordate, cordate- 
orbiculate, or subreniform, strongly 3-lobed, cleft, or parted, 
rarely divided, the segments further shallow-lobed or the 
blades even 5-cleft, the margins dentate-lobed and acumi- 
nate, overall the blades appear regularly laciniate; capitu- 
lescence of solitary heads borne on weak peduncles 3-8 (15) 
em long; heads radiate, cernuous, 10-12 mm high, (5) 7-9 
mm wide, involucres cylindroidal to campanulate, truncate 
at the base; bracts 5.0-9.5 mm long, linear-lanceolate to 
lanceolate-ovate, subattenuate, often purplish; ray flowers 
8-12, ligules yellow, often turning purplish when dry, 2.5- 
9.0 mm long, oblong-elliptic; disc flowers numerous, corollas 
yellow, often purple-tinged, 3.8-5.0 mm long, throats tubu- 
lar-funnelform; achenes (3) 3.5-5.0 mm long, narrowly 
oblong to oblong-elliptic, with prominent callous margins, 
the margins densely ciliate, faces glabrous to puberulent 
at the center; pappus of a crown of squamellae, 0.7-1.0 mm 
lonz, and a single bristle 3.5-6.0 mm long, the bristle very 
stout at the base, tapering to a point, densely barbellate ; 
chromosome number, n = 17. 

Restricted to Baja California Sur in the area of Sierra 
de la Giganta. Flowering year around. (Fig. 5). 

296 Rhodora [Vol. 76 


MEXICO: BAJA CALIFORNIA SUR. Arroyo Carrizal, E of Rancho El 
Horno, NE of San Javier, Carter and Ferris 3811 (UC, US) ; Arroyo 
Santo Domingo, Rancho El Horno, Carter and Ferris 3867 (DS, SD, 
UC); vicinity of Portezuelo E of La Victoria, Carter and Ferris 
3911 (ns, UC); NW of Rancho El Horno, on steep escarpment of 
Mesa de San Alejo, W of San Javier, Carter and Sharsmith 4199 
(Uc); Purisima, Gentry 4213 (ARIZ, Ds, GH, UC); Puerto Escondido, 
Johnston 4115 (CAS, GH, NY, UC, US); ca. 10 mi W of Loreto, Powell 
and Sikes 1663 (SRSC, TEX); ca. 10 mi S of Mission Dolores, Wiggins, 
Carter, Ernst 287 (ps, vs); arroyo Quisapol, E of La Presa, trail 
to Laguna Caquihui, Sierra de la Giganta, Wiggins 15544 (ps). 

Perityle lobata has perhaps been separated from main- 
land Mexico and its relatives there for about four million 
years. At least according to Moore and Buffington (1968), 
this is how long the Baja California peninsula has been 
drifting westward from the mainland. Perityle lobata is 
clearly related to P. palmeri. Distributional considerations 
suggest that the two present-day taxa once had a common 
range (Fig. 5), or else proximal distributions before the 
Sea of Cortez became a barrier of its present width. 

It is possible that Perityle lobata has been introduced to 
the mainland and has hybridized there, accounting for some 
of the variability in mainland taxa such as P. palmeri, In 
fact, it might be postulated that P. palmeri has evolved 
from P. lobata instead of the other way around as previ- 
ously assumed. However, if the latter conjecture were ac- 
curate, one might expect that P. palmeri would be more 
like P. lobata of Baja California. I have not found any 
evidence that P. lobata has been introduced recently to the 

As opposed to Perityle palmeri, P. lobata can be recog- 
nized by its leaf architecture, its purplish stems, pubescence, 
ligule length and number, bract width, achene length, and 
its stout bristles. 

25. Perityle palmeri S. Wats. Proc, Amer. Acad. 24: 57. 
1889. TYPE: Mexico: SONORA: Guaymas, Oct., 1887, E. 
Palmer 308 (Holotype, aH !; isotypes, NY!, uc!, us (3) !). 

1974] Perityle — Powell 297 

Leptopharynx palmeri (S. Wats.) Rydb. N. Amer. Fl. 34: 
23. 1914. 

Plants fleshy-rooted perennials, low and spreading, semi- 
erect, or semipendulent, 10-25 cm high, stems herbaceous 
and rather brittle, often purplish, short or long pilose; 
leaves mostly alternate, short or long pilose, grayish-green, 
6-10 em long, 3.0-5.5 cm wide, cordate, cordate-ovate, to 
cordate-orbiculate, the margins acutely shallow-lobed and 
serrate-dentate, the lobes several, or the blades strongly 
3-lobed, overall the blades may appear irregularly laciniate; 
capitulescence of solitary heads borne on peduncles 2-6 (8) 
cm long; heads radiate, often cernuous at maturity, 8-11 
mm high, 6-13 mm wide, involucres hemispherical to cam- 
panulate and truncate at the base; bracts 7-10 mm long, 
linear to linear-lanceolate and attenuate; ray flowers nu- 
merous, ligules yellow, 7-8 (10) mm long, oblong-elliptic ; 
disc flowers numerous, corollas yellow, 4-6 mm long, throats 
tubular; achenes 2.5-3.5 mm long, narrowly oblong, with 
prominent callous margins, the margins densely long-ciliate, 
faces glabrous or puberulent; pappus of a crown of squa- 
mellae, 0.5-0.9 mm long, and a single bristle, 3-5 mm long, 
the bristle stout, tapering from base to apex, and densely 
barbellate; chromosome number, n = 17. 

Primarily, if not entirely, soil-dwelling, most common in 
the vicinity of Guaymas, Sonora, but also in northern Sina- 
loa, Mexico. Flowering probably year around. (Fig. 5). 


Mexico: SINALOA: Topolobampo, Blakey (ARIZ.) SONORA: Guay- 
mas, Brandegee (POM, UC); Plam Canyon, W side of Sierra Libre, 
35 mi S of Hermosillo, Carter (ARIZ); San Pedro Bay, Craig 691 
(POM); Guaymas, Dawson 1003 (ps, US); cliffs, E side of Guaymas, 
Drouet and Richards 3846 (ps); 2.8 mi N of Guaymas, Lewis 5334 
(sMU); 36 mi S of Hermosillo, Lindsay 1125 (ps); near Guaymas, 
Palmer 1218 (us); Sahuaral Bay, Powell and Averett 1523 (SRSC, 
TEX); 1 mi E of San Carlos Resort, Powell and Sikes 1687 (SRSC, 
TEX); 6 km N of Empalme, Ripley 14307 (CAS); 1 mi E of San 
Carlos Resort, Sikes and Babcock 169 (sRSC, TEX); 1 mi N of Naval 
Base, Guaymas Bay, Sikes and Babcock 176 (SRSC, TEX). 

298 Rhodora [Vol. 76 

Perityle palmeri is best distinguished from P. lobata, 
perhaps its closest relative, and from P. cordifolia by the 
following characters: fleshy roots, weak perennial habit in 
soils, lobing of the leaves, ligule length, bract length, and 
pilose pubescence. 

26. Perityle californica Benth, Bot. Sulph. 23. 1844. 
TYPE: Mexico: BAJA CALIFORNIA: Magdalena Bay, 1837?, 
R. B. Hinds (K!). 

Perityle plumigera Harv. & Gray, Mem. Amer. Acad, II. 
4: 77. 1849. TYPE: Mexico: SONORA: “California” (vicinity 
of Hermosillo), Coulter 279 (Holotype, GH!; fragment, 
uc!, us!). 

Perityle deltoidea S. Wats. Proc. Amer. Acad. 24: 57. 
1889. TYPE: Mexico: BAJA CALIFORNIA: Los Angeles Bay, 
1887, E. Palmer 568 (Holotype, GH!; isotypes, NY!, UC! 

Plants delicate or robust annuals, 6-35 cm high, erect or 
decumbent, sparsely or densely leaved; leaves opposite or 
alternate, glandular-puberulent to short pilose, 1-6 (10) em 
long, 0.5-2.5 (6.0) cm wide, often purplish when mature, 
deltoid, ovate, or cordate, 3-lobed to subcruciform with the 
segments serrate-crenate or shallow-lobed, or the margins 
merely shallow-lobed to serrate; petioles 0.3-1.5 (4.0) em 
long; capitulescence of 1-5 heads borne on delicate pe- 
duncles; heads radiate, 3.5-7.5 mm high, 3.0-5.0 (7.5) mm 
wide, involucres narrowly to broadly campanulate; bracts 
linear, oblanceolate, or obovate; ray flowers 10-15, ligules 
yellow, oblong-elliptic; disc corollas yellow, 2.0-3.0 (3.5) mm 
long, throats tubular to subfunnelform; achenes 1.5-2.5 
(3.0) mm long, narrowly obovate, oblanceolate, to oblong- 
oblanceolate, with thin or prominent callous margins, the 
margins short or long ciliate, faces puberulent to densely 
so; pappus of a crown of squamellae and a single bristle, 
1.5-3.0 (3.5) mm long, the bristle barbellate to subplumose; 
chromosome numbers, n — 13, 12, 11. 

Common in desert soils, Sonora and Sinaloa of mainland 

1974] Perityle — Powell 299 

Mexico, and along most of the Baja California peninsula to 
just S of La Paz. Flowering Dec.-June. (Fig. 6). 


MEXICO: BAJA CALIFORNIA. San Juan Mine, Moran 8117 (DS, Sb, 
uc); Los Angeles Bay, Palmer 568 (GH, NY, UC, US); 4 mi S of 
Rancho Rosarito, Powell and Turner 1808 (sRSC, TEX); El Arco, N 
end of town, Sikes and Babcock 297 (SRSC, TEX); mesa S of Rancho 
Mesquital, Wiggins 11326 (DS, GH, UC). BAJA CALIFORNIA SUR. Isla 
Magdalena, Brandegee s.n. (GH, UC, US); 20 km E of San Ignacio, 
Carter, Alexander, and Kellogg 1972 (ps, UC, US); 26 km NW of 
San Ignacio, Carter, Alexander, and Kellogg 2521 (DS, GH, UC, US); 
Arroyo del Cajon de Tecomaja, SW of Puerto Escondido, Carter and 
Kellogg 2906 (uc); Rancho El Horno, NE of San Javier, Carter 
and Ferris 3789 (uc); SE of La Soledad, Carter 5418 (uc); Arroyo, 
4 mi above La Purisima, Constance 3146 (DS, GH, LL, US); Isla San 
Jose, Moran 3785 (DS, SD, UC); Isla Santa Catalina, Moran 3867 (DS, 
sp); Isla San Marcos, Moran 8983 (sp, US); Isla. Carmen, Moran 
9133 (sp); Isla Danzante, Moran 9253 (sp); Santa Rosalia, Palmer 
185 (CAS,GH, US); ca. 19 mi N of La Paz, Porter 414 (CAS, Ds, Uc); 
20 mi S of El Arco, Powell and Sikes 1650 (SRSC, TEX); 0.5 mi S of 
San Ignacio, Powell and Sikes 1653 (SRSC, TEX); ca. 8 mi S of Santa 
Rosalia, Powell and Sikes 1657 (SRSC, TEX); 33 mi S of Santa Rosalia, 
Powell and Sikes 1659 (SRSC, TEX); Bahia. de la Concepcion, Powell 
and Sikes 1661 (sRSC, TEX); 10 mi W of Loreto, Powell and Sikes 
1664 (SRSC, TEX); 13.6 mi S of Mulege, Powell and Turner 1835 
(SRSC, TEX); 11.5 mi W of Loreto, Powell and Turner 1837 (SRSC, 
TEX); 15 mi W of San Javier, Powell and Turner 1845 (SRSC, TEX) ; 
3 mi E of La Paz, Sikes and Babcock 216 (SRSC, TEX); near km 
40 on hwy between La Paz and Santo Domingo, Thomas 8455 (CAS) ; 
Isla Partida, Wiggins, Carter, and Ernst 412 (DS); Commondu, 
Wiggins 5502 (CAS, DS, GH, NY, UC, US); 22 mi N of San Ignacio, 
Wiggins 7888 (DS, GH, TEX, UC, US); 20 mi S of Mulege, Wiggins 
11408 (CAs, ps, GH, UC, US); 6 mi W of Canipole, Wiggins 11489 
(CAS, DS, GH, Uc, US); Llano Caquihui, W of Los Dolores, Wiggins 
15516 (ps, US); 16 mi E of La Paz, Wiggins 15625 (Ds, us); Isla 
Carmen, Wiggins 17497 (ps). SINALOA. N of Topolobampo, Flyr 72 
(TEX); vicinity of San Blas, Rose, Standley, and Russell 13244 
(NY, US); vicinity of Fuerte, Rose, Standley and Russell 13472 (NY, 
us); W side of Topolobampo Harbor, Sikes and Babcock 189 (SRSC, 
TEX). SONORA. 8 mi N of Hermosillo, Breedlove 1758 (ns); San 
Pedro Bay, Craig 662 (PoM); E side of Guaymas, Drouet and 
Richards 3844 (CAS, DS, GH, US); 73 mi S of Hermosillo, Flyr 98A 
(TEX); 47 mi N of Hermosillo, Frye and Frye 2305 (GH, us); 1 mi 
E of San Carlos Resort, Powell and Sikes 1686 (SRSC, TEX); 1-2 mi 

300 Rhodora [Vol. 76 

Fig. 6. Distribution of P. californica (closed circles); the bars 
indicate populations from which chromosome numbers were taken, 
with (1) n — 11 II, (2) n = 12 II, (3) n — 13 II, and (4) n —10 
II + 1 III; all mainland counts have been n= 13. P. cuneata var. 
cuneata (open circles) ; P. cuneata. var. marginata, (closed triangles). 

1974] Perityle — Powell 301 

N of San Carlos Resort, Powell and Sikes 1691 (SRSC, TEX) ; vicinity 
of Navojoa, Rose, Standley, and Russell 13140 (Ny, US); 110 mi 
E of Hermosillo, Sikes and Babcock 151 (SRSC, TEX); 32 mi N of 
Hermosillo, Shreve 10053 (LL); 10 mi N of Hermosillo, Wiggins 6250 
(ns, POM, US); 5 mi SE of Torres, Wiggins 6273 (DS, GH, POM, US). 

In her discussion of P. californica, Everly (1947) em- 
phasized the variable habit of this taxon. The species oc- 
curs in Baja California and in Sinaloa-Sonora of mainland 
Mexico. Most of its morphological variability is expressed 
in plant size, number of leaves, and leaf shape. Floral char- 
acteristics are essentially the same throughout the distribu- 
tional range. 

Chromosomal variability also exists in Perityle californica 
where numbers of n — 13, 12, 11, and 10 II + 1 III have 
been reported (Powell, 1968; 1972b). Some distributional 
and morphological correlation with chromosome number 
has been observed, although no taxonomic status is war- 
ranted for the cytotypes. Plants of the mainland popula- 
tions are n = 13, as so far determined, and tend to be 
rather tall and robust with deeply lobed (often trilobed) 
leaves. In wet years P. californica is a common weed in 
some parts of Sonora and Sinaloa. Chromosome numbers 
of n = 13, 12, 11, and 10 II + 1 III have been found in 
Baja California populations (Fig. 6), and the plants are 
usually small with leaves not deeply lobed (mostly ser- 
rately-lobed). I have found the Baja California populations 
to be invariably small and scattered, but I have not seen 
the peninsula in a wet year. Populations with » — 11 are 
most common in Baja California and probably extend 
throughout the range of the species (Fig. 6). One popula- 
tion with n = 12 has been found, and those with n = 10 
II + III presumably arose through hybridization of cyto- 
types with n = 12 and n = 11 (Fig. 6). The one peninsula 
collection with » — 13 (Fig. 6) could have been introduced 
from the mainland, or seemingly less likely at present, could 
represent the presence of native n — 13's on the peninsula. 
It seem reasonable to conclude that P. californica in Baja 
California evolved by aneuploid reduction in chromosome 
number from mainland cytotypes. 

302 Rhodora [Vol. 76 

Perityle californica does not share close or obvious rela- 
tionship with any species, and its origin as a low-numbered 
aneuploid is not clear. The species is superficially similar 
to P. cuneata (n = 16, 12), but these taxa differ consider- 
ably in floral characters, particularly of the achenes and 
pappus. Indeed, the floral] differences between these two 
aneuploid species are significant enough to cast considerable 
doubt upon their common origin. Perityle californica has 
narrow oblanceolate achenes with a single, long, subplumose 
pappus bristle, while P. cuneata has broad, obcordate or 
obovate achenes with two delicate pappus bristles. The 
morphology of P. californica (including flower color and 
achene and pappus features) resembles that of the P. cordi- 
folia group (x = 17), and possibly originated therefrom by 
aneuploid reduction. 

Certain aspects of the Perityle californica synonymy are 
somewhat confusing, but a good explanation of the histori- 
cal situations was given by Everly (1947). 

27. Perityle cuneata Brandeg. Zoe 1: 54. 1890. TYPE: 
Mexico: BAJA CALIFORNIA: Sierra de Laguna, near Todos 
Santos, 21 Jan. 1890, T. S. Brandegee (Holotype, UC!; iso- 
type, US!). 

27a. Perityle cuneata var. cuneata. 

Plants robust annuals, 25-50 (80) cm high, erect or de- 
cumbent, sparsely to densely leaved; leaves opposite or 
alternate, glandular-puberulent to nearly glabrous, 3-8 (15) 
em long, 1.5-4.5 (11.0) cm wide, often purplish when ma- 
ture, cordate to broadly so, ovate, or subhastate, the mar- 
gins serrate-lobed to serrate-crenate; petioles 0.6-2.0 (5.0) 
em long; capitulescence of 1-3 heads borne on rather stout 
peduncles; heads radiate, 5-8 (10) mm high, 6-10 (12) mm 
wide, involucres campanulate to hemispherical; bracts ob- 
lanceolate to obovate and attenuate; ray flowers 10-15, 
ligules yellow, oblong-elliptie; disc corollas yellow, 3.0-3.5 
mm long, throats tubular; achenes (2.5) 3.0-3.8 mm long, 
obcordate-cuneate to oblanceolate, with broad callous mar- 

1974] Perityle — Powell 303 

gins, often purplish, the margins puberulent, faces glabrous 
to puberulent; pappus of a crown of squamellae and 2 
(rarely 0-2) delicate bristles, 0.5-1.2 (2.0) mm long; chro- 
mosome number, n = 12. 

Infrequent in soils, Cape Region of Baja California Sur 
to just S of La Paz, most common on eastern side. Fower- 

ing Jan.-Sept. (Fig. 6). 


Mexico: BAJA CALIFORNIA SUR: San Jose del Cabo, Brandegee (DS, 
GH, US) ; Sierra San Francisquito and La Chuparosa, E side of Sierra 
de la Victoria, Carter and Ferris 3386 (DS, SD, UC); ca. 3 km above 
Boca de la Sierra, Moran 7091 (CAS, DS, GH, SD, TEX); Potrero de 
Almenta, S fork of Canyon San Pedro, Moran 7387 (CAS, DS, SD) ; 
ca. 5 mi S of San Antonio, Powell and Sikes 1667 (SRSC, TEX) ; Boca de 
la Sierra, Powell and Sikes 1671 (sRSC, TEX) ; 2 mi S of San Antonio, 
Sikes and Babcock 233 (SRSC, TEX); ca. 2 mi N of San Antonio, 
Sikes and Babcock 265 (sRSC, TEX); ca. 6 mi SW from Santiago, 
Thomas 7715 (CAS, DS, SD, US); ca. 5 mi SW of La Palma and ca. 
8 mi NW from Santa Anita, Thomas 7738 (DS, SD, UC). 

The two varieties of Perityle cuneata are morphologi- 
cally similar, although extreme forms of these taxa do ex- 
hibit considerable differences. Everly (1947) recognized 
P. cuneata and P. cuneata, var. marginata as conspecific. 
In addition to the morphological differences, I have recog- 
nized var. cuneata, (n = 12) and var. marginata (n = 16) 
because of their chromosome numbers, Indeed, the chromo- 
somal differences suggest that further study might demand 
the elevation of var. marginata to specific rank. 

Besides its chromosome number, Perityle cuneata var. 
cuneata is characterized by robust habit, large cordate 
leaves, large heads, tubular disc corollas, obcordate-cuneate 
achenes, and attenuate involucral bracts. The plants of var. 
marginata are usually smaller with smaller leaves, smaller 
heads and other floral characters, campanulate-funnelform 
dise corollas, obovate achenes, and acute involucral bracts. 
In addition, var. cuneata usually occurs in higher hills and 
canyons of the Cape Region while var. marginata occurs 
mostly in sandy places not far above sea level. 

304 Rhodora [Vol. 76 

Perityle cuneata does not exhibit a close morphological 
relationship with any other species. Superficially, P. cu- 
neata most resembles P. californica, but it possibly evolved 
independently of the latter taxon by aneuploid reduction 
from the P. cordifolia line. 

27b. Perityle cuneata var. marginata (Rydb.) I. M. Johns- 
ton, Proc. Calif. Acad. IV, 12: 1204. 1924. 

Perityle marginata Rydb. N. Amer. Fl. 34: 14. 1914. 
TYPE: Mexico: BAJA CALIFORNIA: San Jose del Cabo, 25 
Mar. 1911, J. N. Rose 16459 (Holotype, NY!; fragment, 
UC!; isotypes, GH !, US!). 

Plants delicate or robust annuals, 6-40 cm high, erect or 
decumbent; leaves 1.5-5.0 (10) em long, 0.9-3.0 (5.0) cm 
wide, cordate, ovate, or deltoid, the margins doubly serrate- 
crenate to shallow-lobed; petioles 0.2-1.8 (4.0) em long; 
heads radiate, 4-7 mm high, 4-7 mm wide; bracts subovate, 
oblanceolate, or linear-lanceolate and acute; ligules yellow; 
dise corollas yellow, 2 (2.5) mm long, throats narrowly 
campanulate, campanulate-funnelform, rarely subtubular; 
achenes 1.5-2.0 (2.5) mm long, obovate, subcuneate, rarely 
almost oblong, with prominent callous margins, the margins 
short or long-ciliate; pappus of a crown of squamellae and 
2 (rarely none) delicate bristles, 0.5-1.0 (1.5) mm long; 
chromosome number, n = 16. 

Infrequent in desert soils, Cape Region of Baja California 
Sur and N to near La Paz, most common on southern and 
western sides. Flowering probably year around. (Fig. 6). 


Mexico: BAJA CALIFORNIA SUR. San Jose del Cabo, Jones s.n. (NY, 
POM, SD, UC); Arroyo de los Pozos, Moran 6886 (CAS, DS, GH, TEX) ; 
2.5 km NE of Cabo San Lucus, Moran 7049 (CAS, DS, GH, SD) ; 6.4 mi 
N of Todos Santos, Porter 79 (CAS, DS, UC); 1.4 mi W of El Coyote, 
Porter 118 (CAS, DS, UC); ca. 17 mi S of San Antonio, Powell and 
Sikes 1668 (SRSC, TEX); ca. 9 mi S of San Bartolo, Powell and Sikes 
1670 (SRSC, TEX); 26 mi NE of Todos Santos, Powell and Sikes 1680 
(SRSC, TEX); 2 mi NE of Eureka, Powell and Turner 1848 (SRSC, 
TEX); ca. 1 mi N of Pescadero, Sikes and Babcock 255 (SRSC, TEX) ; 
near km 40 between La Paz and Santo Domingo, Thomas 8455 (Ds, 

1974] Perityle — Powell 305 

GH); 5 mi W of San Jose del Cabo, Wiggins 14370 (CAS, DS, TEX, UC) ; 
8 mi E of Cabo San Lucas, Wiggins 14645 (CAS, DS, GH, TEX, UC). 

The NY (holotype) and UC specimens of the type collec- 
tion (Rose 16459) are typical of var. marginata, while the 
US and GH specimens (isotypes) approach var. cuneata. It 
is possible that the type collection may be mixed, containing 
plants of both varieties. 

Some of the small forms of var. marginata are so like P. 
californica superficially that the two taxa can not be told 
apart except by microscopic examination of the achenes 
and pappus. The Thomas 8455 collection includes plants 
of both var. marginata and P. californica, but I have not 
otherwise noted sympatric occurrence of the taxa. No 
interspecific hybridization has been detected. 


ALSTON, R. E., & B. L. TURNER. 1963. Natural hybridization among 
four species of Baptisia (Leguminosae). Am. Jour. Bot. 50: 

EvERLY, M. L. 1947. A taxonomic study of the genus Perityle and 
related genera. Contrib. Dudley Herb. 3: 375-396. 

JOHNSTON, I. M. 1929. In: Papers on the Flora of Northern Chile. 
Contrib. Gray Herb. 85: 127-128. 

Moore, D. G., & E. C. BUFFINGTON. 1968. Transform faulting and 
growth of the Gulf of California since the late Pliocene. Science 
161: 1238-1241. 

NiLES, W. E. 1970. Taxonomic investigations in the genera Perityle 
and Laphamia (Compositae). Mem. N. Y. Bot. Gard. 21: 1-82. 

PowELL, A. M. 1968a. Additional discussions pertaining to the con- 
generic status of Perityle and Laphamia (Compositae). Sida 3: 

1968b. Chromosome numbers in Perityle and related 
genera (Peritylinae — Compositae). Am. Jour. Bot. 55: 820-828. 
1969. Taxonomy of Perityle section Pappothrix 
(Compositae — Peritylinae). Rhodora 71: 58-93. 

. 1970. Natural intersectional hybridization in Peri- 
tyle (Compositae). Brittonia 22: 3-10. 

1972a. Taxonomy of Amauria (Compositae — Peri- 
tylinae). Madroño 21: 516-525. 

1972b. Peritylinae, In IOPB Chromosome Number 
Reports. Taxon 21: 344. 

306 Rhodora [Vol. 76 

1972c. Artificial hybridizations in the subtribe Peri- 

tylinae (Compositae — Helenieae). Am. Jour. Bot. 59: 760-768. 

1972d. Taxonomy of Pericome (Compositae — Peri- 

tylinae). Southwest. Nat. 18: 335-339. 

1972e. New species of Perityle and Amauria (Com- 

positae). Madroño 21: 456-458. 

1973. Taxonomy of  Perityle section Laphamia 

(Compositae — Peritylinae). Sida 5: 61-128. 

, & S. Sikes. 1970. Chromosome numbers of some 

Chihuahuan Desert Compositae. Southwest. Nat. 15: 175-186. 

,& H. S. Tsang. 1966. Preliminary chromatographic 
studies of Laphamia and related genera of Compositae. South- 
west. Nat. 11: 190-195. 

RAVEN, P. H. 1963. Amphitropical relationships in the floras of 
North and South America. Quart. Rev. Biol. 38: 151-177. 

RYDBERG, P. A. 1914. Carduaceae — Helenieae — Peritylinae. N. 
Am. Fl. 34: 11-27. 

SHINNERS, L. H. 1959. Species of Laphamia transferred to Perityle 
(Compositae — Helenieae). Southwest. Nat. 4: 204-209. 

Penduletin 4'-0-Methyl Ether, a new flavonoid from Perityle 
vaseyi (Compositae). Phytochemistry 11: 2351. 

STEBBINS, G. L. 1950. Variation and evolution in plants. Columbia 
Univ. Press, New York. 

TURNER, B. L. 1966. Taxonomy of Eutetras (Compositae — Heleni- 
eae). Southwest. Nat. 11: 118-122. 

TURNER, B. L. & D. FLYR. 1966. Chromosome numbers in the Com- 
positae. X. North American species, Am. Jour. Bot. 53: 24-33. 


HAMPSHIRE. South Bay Bog at the southern end of First 
Connecticut Lake in Pittsburg, Coos County, New Hamp- 
shire apparently was not visited by A. S. Pease, S. K. 
Harris and other botanists who collected extensively in 
Coós County. The bog is very extensive, much of it open 
but with scattered clumps of stunted Black Spruce and 
Tamarack. Lying between 1600 and 1700 feet elevation, 
it appears in every way to be boreal in character with 
the following species of shrubs in abundance, Ledwm 
groenlandicum, Rhododendron canadense, Andromeda glau- 
cophylla, Kalmia polifolia, K. angustifolia, Cassandra. caly- 
culata, Pyrus melanocarpa, Viburnum cassinoides and 
Vaccinium angustifolium. 

Bog Branch of Cedar Stream flows through or along the 
northern edge of South Bay Bog and near it, in sites where 
some drainage appears to run off the adjacent upland, there 
are scattered trees of Thuja occidentalis L., some up to 
30 inches d.b.h. These represent remnants of the primeval 
forest since, along with dying trees of large size, there are 
fallen ones in all stages of decay. Several species of orchids 
grew in the immediate vicinity of trees of Arbor Vitae, the 
rarest of which was Listera convallarioides (Liv.) Nutt. 
A plant here of perhaps greater interest was Epilobium 
palustre L. var. palustre of which Seymour (1969) states 
“Often with Thuja,” but of which he cites only one locality 
in New Hampshire, “Mt. Washington." Pease (1964) states 
its habitat as “Boggy places, especially on the high moun- 
tains,” but he gives two localities for it at lower elevations 
both probably with some “Cedar” associated, “Stewarts- 
town, Cedar bog” and “Columbia, near Lime Pond.” 

In the more open part of South Bay Bog dominated by 
sphagnum moss and the previously mentioned shrubs and 
dwarf conifers there are plants of still greater interest to a 
taxonomist, one being Carex exilis Dewey and the other 
Geocaulon lividum (Richards.) Fern. Carex exilis is a sedge 
of local but wide distribution in New England, it being 


308 Rhodora [Vol. 76 

found occasionally in all States except Connecticut. Fer- 
nald (1950) omits mention of New Hampshire and Vermont 
in his statement of range for the species though he does 
mention Massachusetts, while Seymour (1969) gives but 
one township for it in New Hampshire, “Tamworth” where 
it was collected by F. L. Steele in a bog near Chocorua 
Lake and where he has shown it to me. Specimens of it are 
in the herbaria of the New England Botanical Club (NEBC) 
and the University of New Hampshire (NHA). However 
also at NHA are two other collections of fairly long stand- 
ing, the earlier being from Windham in Rockingham County 
collected on June 14, 1897 by W. S. Harris. This record 
was included in the list published by Hodgdon and Fried- 
lander (1949). The second collection of Carex exilis at 
NHA is from a bog in Pittsburg known as “Moose Pasture” 
lying along East Inlet about two miles northeast of Second 
Connecticut Lake. This was collected by me in company with 
Peter Allen on July 23, 1960 and bears my collection num- 
ber 11636. Pease (1964) did not include C. exilis in his 
"Flora" and it is quite possible that this specimen of it 
was not available for inspection at the time that Pease 
was gleaning additional records for Coós County and Harris 
was carefully checking the manuscript before publication. 
The specimens of C. exilis observed at South Bay Bog on 
our visit of June 8, 1973 were completely dioecious, the 
clones or colonies of female individuals being widely 
separated from the males. 

The other species of outstanding interest at South Bay 
Bog was Geocaulon lividum. I have been quite familiar 
with the Northern Comandra from the frequency of its 
occurrence in bogs on Campobello Island, New Brunswick 
and its presence on the Wolf Islands in the Bay of Fundy 
where R. B. Pike and I have observed it. It seems strange 
that a species inhabiting bogs at low elevations in coastal 
eastern Maine and the Maritime Provinces should be con- 
fined to montane areas inland though the same relationship 
seems to hold for Rubus chamaemorus L. and Lycopodium 
annotinum L. var. pungens (LaPyliae) Desv. Harris (1965) 

1974] Pittsburg, N. H. Plants — Hodgdon 309 

discussed the distribution of Geocaulon lividum in a critical 
way, pointing out that its habitat in stations that he visited 
was mostly damp but not boggy sites and that the lowest 
elevation for it was at 2500 ft. on the Basin Rim in Coos 
County. The station at South Bay Bog is in no way mon- 
tane and the elevation is between 1600 and 1700 feet. It is 
entirely possible that botanists have concentrated their 
collecting more in the mountains than in the more prosaic 
terrain of bogs that often lie between the mountains, thus 
passing by an interesting intermediate type of habitat for 
certain species that are supposed to occur only in coastal 
areas or near the tops of mountains. In any event it is 
evident that Geocaulon lividum should be searched for 
widely as Stuart Harris pointed out, not only as he sug- 
gested, in montane situations but also in bogs at inter- 
mediate elevations in Northern New England. 

Specimens of Carex exilis and Geocaulon lividum have 
been deposited in the herbaria of the New England Bo- 
tanical Club (NEBC) and the University of New Hampshire 

These discoveries would not have been possible except 
for the expert guidance of Dr. Peter Allen. In addition 
I have borrowed freely in preparing this article from notes 
that he made or that we recorded jointly on the occasion 
of our visit to South Bay Bog on June 8, 1973. 


FERNALD, M. L. 1950. Gray’s Manual of Botany, eighth edition, 
American Book Company, N. Y. 1682 p. 

Harris, S. K. 1965. Geocaulon lividum in New England. Rhodora 
67: 407-409. 

HopGDoN, A. R. and HERBERT FRIEDLANDER. 1949. Additions to the 
Harris “Flora of Windham, New Hampshire.” Rhodora 51: 110. 

PEASE, A. S. 1964. A Flora of Northern New Hampshire, New 
England Botanical Club Inc. Cambridge, Mass. 278 p. 

SEYMOUR, F. C. 1969. The Flora of New England. The Charles 
Tuttle Co., Rutland, Vermont. 596 p. 


DURHAM, NEW HAMPSHIRE. Pittsfield, N. H. (Mer- 
rimack County) is no longer the southernmost collection 
point in New Hampshire for Nuphar microphyllum (Pers.) 
Fern. as reported in my article *Some Morphological Aids 
in Distinguishing Nuphar microphyllum from Similar 
Aquatics” (Rhodora 75: 65-74, March 1973). On October 8, 
1973 while A. R. Hodgdon and I were scouting the Lamprey 
River in Durham as a possible field trip site for his course 
in aquatic plants at the University of New Hampshire, 
I discovered a colony of N. microphyllum growing with 
N. variegatum and Nymphoides cordata in about two feet 
of water in a small cove near the upper end of Moat Island. 
Perhaps closer observation may reveal some plants of the 
hybrid, Nuphar X rubrodiscum. Undoubtedly other sta- 
tions for N. microphyllum exist but its relative rarity in 
southern New Hampshire has made it unnoticeable to the 
casual observer. The two specimens collected late in the 
season are a bit ragged and will be kept as vouchers in 
the NHA Herbarium. An attempt will be made to secure 
more presentable material for the New England Botanical 
Club Herbarium. 

DURHAM, N. H. 03824 



The type of Chamaesaracha coronopus Was erroneously 
listed as Berlandier 1494 (= 234) (Rhodora 75: 339). 
The section referring to the type material should read: 

Chamaesaracha coronopus (Dunal) Gray, Bot. Calif. 1: 
540. 1876. Holotype (G!), [Texas, Bexar Co.], Rio 
Medina, between Laredo and Bejar [San Antonio]. 1828. 
Berlandier 1513 (= 253). Isotypes: GH! & K!. 

The locality indicated on the holotype and original de- 
scription is only between Laredo and San Antonio, then 
Bejar, Mexico. An isotype at Kew, however, has Rio 
Medina on the label. The Medina River flows southeast 
from Bandera County into Medina County, then east into 
Bexar County. The route taken by Berlandier likely 
erossed the Medina River in Bexar County, about 12 miles 
south of San Antonio. 


LATA. While I was walking through a swampy area close 
to Spruce Hole Bog in Durham, N.H. on October 20, 1973, 
a fern frond came to my attention that from afar resembled 
an aberrant form of sensitive fern, Onoclea sensibilis. Upon 
further investigation, however, what was found turned out 
to be the fertile frond of Woodwardia areolata (L.) Moore, 
the netted-chain fern, so named for its long sori arranged 
in a row on each side of the secondary midrib. 

The discovery of this fern is noteworthy when one real- 
izes that only two stations for it have ever been recorded 
from New Hampshire, both in Rockingham County: Rye 
Beach by Walter Deane in 1886 (N.E.B.C.) and Seabrook 
by A. A. Eaton in 1896 (N.E.B.C.) In the seventy-seven 
years since those recorded findings, Woodwardia areolata 
has not been found in New Hampshire. In addition, this 
new station extends the range of this coastal plain species 
by approximately twenty miles. 

Shaded by two bushes of Vaccinium corymbosum, a total 
of six plants have been located within an area of three feet. 
The plants are growing on rotting stump matter and are 
surrounded most of the year by a foot of water or more. 

Such a find always leads one to speculate as to how 
this fern came to occur at this location, how long it has 
been there, and why it was not discovered long before this. 

A voucher specimen of the Woodwardia areolata is de- 
posited in the University of New Hampshire herbarium. 





With this number the present editor-in-chief with twelve 
and a half years of service in that capacity, is completing 
his tenure. Also Dr. Radcliffe B. Pike, as a valued associate 
editor during this entire period, is completing his editorial 

We trust that the younger group of editors who are now 
in charge will find the journal to be prospering. Many 
changes in Rhodora have been made during the past twelve 
years; we express the hope that the journal can continue 
to be molded to fit changing needs. 

The new editor-in-chief is Dr. Alfred Linn Bogle, 
Associate Professor of Botany at the University of New 
Hampshire. Please send all manuscripts and communica- 
tions to Dr. A. Linn Bogle, Department of Botany and 
Plant Pathology, University of New Hampshire, Durham, 
New Hampshire 03824. 





Manuscripts must be double-spaced or preferably triple- 
spaced (not on corrasable bond), and a list of legends for 
figures and maps provided on a separate page. Footnotes 
should be used sparingly, as they are usually not necessary. 
Do not indicate the style of type through the use of capitals 
or underscoring, partieularly in the citations of specimens, 
except that the names of species and genera may be under- 
lined to indicate italics in discussions. Specimen citations 
should be selected critically especially for common species 
of broad distribution. Systematic revisions and similar 
papers should be prepared in the format of “The System- 
aties and Ecology of Poison-Ivy and the Poison-Oaks," 
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu- 
larly with reference to the indentation of keys and syno- 
nyms. Papers of a floristic nature should follow, as far as 
possible, the format of “Contribution to the Fungus Flora 
of Northeastern North America. V.," H. E. Bigelow & M. E. 
Barr, Rhodora 71: 177-203. 1969. For bibliographie cita- 
tions, a recommended list of standard journal abbreviations 
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. 
Club 85: 277-300. 1958. 

Volume 76, No. s06, including pages 151-314, was issued July 3, 1924. 

CONTENTS: — continued 

New Station for Nuphar microphyllum in Durham, New 

Johonet C. Wicks eese eee eene eene erano peonia ei sinit 310 
Typification of Chamaesaracha coronopus 

John B: Averell .cictcocesdacsncsisccanscnectsiescscavencecusscesepsatstgseesteduccsese 811 
Range Extension of Woodwardia areolata 

Aminta Kit field ............. 4. eee eere eese eese eee esee estan nost tatto a ah taa a 312 
Editorial Announcement ................ eere eene eene nnne ennas 813 

Instructions for Contributors .............. eene 314 




Conducted and published for the Club, by 
ALFRED LINN BOGLE, Editor-in-Chief 


Associate Editors 

Vol. 76 September, 1974 No. 807 
A Systematic Study of the Genus Kalmia (Ericaceae) 

TORN. E DIIP aoieanna aia E e 315 
Marine Algae of Rhode Island 

R. D. Wood and Martine Villalard-Bohnsack .................. 399 

A Biosystematic Revision of Lupinus lyallii 
20) 08 OT es Ts canioniusnndctsvsensereaseesanssise A E 422 

The Role of Some Haloragaceae in Algal Ecology 
L, C. Colt, Jr., and C. Barre Hellquist ................... 446 

Illinois Field and Herbarium Studies 
Robert H. Mohlenbrock and Dan K. Evans ......... 460 

(Continued on Inside Back Cover) 

Che Nef England Botanical Club, Ine. 
Botanical Museum. Oxford St.. Cambridge, Mass. 02138 

RHODORA. — A quarterly journal of botany, devoted primarily to the 
flora of North America and floristically related areas. Price $10.00 
per year, net, postpaid, in funds payable at par in the United States 
currency at Boston. Some back volumes, and single copies are 
available. For information and prices write RHODORA at address 
given below. 

Scientific papers and notes, relating directly or indirectly to the 
plants of North America, will be considered by the editorial com- 
mittee for publication. Articles concerned with systematic botany 
and cytotaxonomy in their broader implications are equally accept- 
able. All manuscripts should be submitted in duplicate, and should 
be double-spaced throughout. Please conform to the style of recent 
issues of the journal. Illustrations can be used only if the cost of 
engraver's blocks is met through the author or his institution. 
Forms may be closed five weeks in advance of publication. Ex- 
tracted reprints, if ordered in advance, will be furnished at cost. 

Address manuscripts and proofs to 

Dept. of Botany, University of New Hampshire, 
Durham, New Hampshire 03824 
Subscriptions and orders for back issues (making all remittances 
payable to RHODORA) should be sent to RHODORA, Botanical 

Museum, Oxford Street. Cambridge, Mass. 02138. In order to receive 

the next number of RHODORA, changes of address must be received 

prior to the first day of March, June, September or December. 
Second Class Postage Paid at Boston, Mass. 




Vol. 76 September, 1974 No. 807 



A study of the cytology and genetics of the genus Kalmia 
was started in 1961 by Dr. Richard A. Jaynes of the Con- 
necticut Agricultural Experiment Station. At that time 
little was known about the genetics of this group although 
a few of the species are important ornamental shrubs. Dr. 
Jaynes has now accumulated many living representatives 
of most of the species due to the efforts of numerous bota- 
nists and friends. As a result of his studies (Jaynes 1968a, 
1968b, 1969, 1971b) information is now available concern- 
ing cross compatibility, speciation, and possible evolution- 
ary relationships within the genus. 

As treated here, the genus is composed of seven species, 
six varieties and numerous forms, both natural and culti- 
vated. The genus is restricted to North America and Cuba, 
being particularly abundant along the east and west coasts 
and at higher elevations and latitudes. No monographic 
study of the genus has been undertaken, but the species 
have been adequately described in many floristic works, and 
Small (1914) treated all of the species that are presently 
recognized. Since one of the species is commonly cultivated 
and all are probably poisonous, this taxon has also been 
reviewed in horticultural and agricultural literature. This 
present study is an attempt to bring together most of this 


316 Rhodora [Vol. 76 

information in the light of what is now know about the 
geneties and cytology of the genus. 


The author is grateful to the many people who helped in 
the completion of this study. Dr. Richard A. Jaynes pro- 
vided access to his herbarium specimens and living material 
as well as his notes and extensive knowledge concerning 
the cytology and genetics of the genus. The Connecticut 
Agricultural Experiment Station provided space and equip- 
ment for 4 months to examine and collect material. Also, 
many individuals at Eastern Illinois University provided 
helpful criticism and advice in the preparation of the manu- 
script. The author is also indebted to the many curators 
who sent specimens for examination. Over 8,000 specimens 
were examined from 75 herbaria from the United States, 
Canada, and Europe. In the lists of representative speci- 
mens these herbaria are indicated by their abbreviations 
following Lanjouw and Stafleu (1964). 


References to mountain laurel, Kalmia latifolia, appear 
in the early literature of the American colonies. The Cape 
Cod “Rose-trees” of Henry Hudson's log (Purchas, 1625) 
may have been this species (Holmes, 1956), and Smith 
(1624) mentions that laurel occurs in Virginia. This spe- 
cies was first described and figured by Plukenet (1696) 
under the name Cistus sempervirens, described by Grono- 
vius (1739) who called it “The Common Laurel/vulgarly 
called Ivy," while the first color print was. figured by 
Catesby (1743) under the name Chamaedaphne. 

Because of their potential as ornamentals, some of the 
species were introduced into Europe. According to Aiton 
(1789) Kalmia latifolia and K. angustifolia were intro- 
duced into England in 1734 and 1736 by Peter Collinson, 
an English merchant and fellow of the English and Swedish 
Royal Societies. A third species, K. polifolia, was intro- 

1974] Genus Kalmia — Ebinger 317 

duced in 1767. Catesby (1743) reported that K. latifolia 
from Pennsylvania flowered in his garden at Fulham in 
1740. Numerous horticultural forms have been developed 
as a result of breeding and selection in Europe for the past 
200 years. 

Probably the first detailed account of the toxic character- 
istics of the genus occur in Peter Kalm's journal (Benson, 
1937) in which he described in detail the poisonous proper- 
ties of the “Laurel Trees" (Kalmia latifolia). He also 
described both K. angustifolia and K. latifolia as to their 
characteristics, economic importance, habitat requirements 
and general distribution. 

The generic name Kalmia was first proposed by Chenon 
(1751) in a doctoral thesis, and was based on specimens 
that Peter Kalm had collected during his trip to the New 
World. Kalmia latifolia and K. angustifolia are described 
in this thesis, and Linnaeus (1753) included both in his 
Species Plantarum. Later, Linnaeus (1754) described the 
genus and placed it in his first order (Monogynia) of his 
tenth class (Decandria). 

The genus is now regarded as a relatively primitive mem- 
ber of the Ericaceae. Drude ( 1889) placed it in the Tribe 
Phyllodoceae Drude of the subfamily Rhododendroideae 
Drude, and its position has not been changed by most 
subsequent authors. The members of this tribe are gen- 
erally evergreen plants, usually with buxoid or ericoid 
leaves, with regular flowers, and with seeds that usually 
lack appendages. Two microscopic characteristics of the 
anthers have also been used to characterize the tribe: the 
presence of resorption tissue involving the epidermis of 
the anther and the absence of filaments among the pollen 
tetrads. Copeland (1943) discussed in detail the morphol- 
ogy and anatomy of the tribe Phyllodoceae and considered 
the genera Rhodothamnus Reichenb., Kalmia, and Phyllo- 
doce Salisb. to be relatively primitive, an opinion also ex- 
pressed by Wood (1961). Studies of wood anatomy by Cox 
(1948) indicate that the genus Kalmia is closely related to 

318 Rhodora [Vol. 76 

Rhodothamnus and that these two genera represent one of 
the three lines of evolution in the Phyllodoceae. 

There have been few nomenclatorial complications in the 
genus, and only two other generic names have been pro- 
posed. Kuntze (1891) used Catesby's pre-Linnean name of 
Chamaedaphne and made wholesale transfers to it, while 
Small (1914) divided Kalmia into two genera. He proposed 
the genus Kalmiella for Kalmia hirsuta and its related 
species that differ by having a deciduous calyx and flowers 
borne singly in the axils of ordinary or somewhat reduced 
leaves. In all other morphological and anatomical character- 
istics these species do not differ from the other members 
of the genus, and there is no real justifieation for their 
removal. Except for Alain (1957), all subsequent authors 
have treated these species as members of the genus Kalmia. 


The genus Kalmia is economically important mainly be- 
cause a few of the species are commonly grown as orna- 
mentals. By far the most important species is the moun- 
tain laurel, Kalmia latifolia, which is commonly planted as 
an ornamental throughout its natural range as well as in 
other parts of North America and Europe where climate 
and soil are satisfactory. This species varies widely in 
flower color, foliage, and size, and many cultivars have been 
named and described (Holmes, 1956; Dudley, 1967). Culti- 
vars of mountain laurel had been named by 1840, and at 
the present time at least 90 names have been applied to 
different forms of the species. This species has been sug- 
gested as a possible choice for the national flower of the 
United States (Van Rensselaer, 1889), and presently it is 
the state flower of Connecticut and Pennsylvania. 

The sheep laurel, Kalmia angustifolia, is also becoming 
an important cultivated plant. Unlike the mountain laurel, 
this species can be propagated by cuttings. A number of 
cultivated forms are available, and selections should become 
available as soon as nurserymen become familiar with hand- 

1974] Genus Kalmia — Ebinger 319 

ling them. This species is not important for its wood as 
few specimens exceed 3 feet in height, but it is economically 
important as a weed. Smith, Hodgdon and Eggert (1947) 
considered it an important weed in low-bush blueberry fields 
and tried fire, continued cutting, and lime to control it. 
Later, Trevett (1956) reported on the effects of 2,4-D and 
24,5-T in controlling sheep laurel in blueberry fields. 
Porcher (1869) found that the leaves and twigs of this 
species will dye cotton a fine drab color if a copper mordant 
is used. 

The foliage of Kalmia latifolia is commonly gathered in 
eastern United States for floral displays as well as for 
Christmas decorations and other festivities, continuing a 
tradition started in colonial days. As early as 1913, Britton 
suggested that this species should probably be protected 
since it was becoming rare in some areas due to indiscrimi- 
nate collecting. Buttrick (1924), in an extensive article 
concerning growth and cultivation of mountain laurel in 
Connecticut, estimated that the amount of foliage of this 
species used in New York City probably exceeded 1,000 
tons per year and that the maximum amount used in the 
United States for decorative purposes probably exceeded 
10,000 tons per year. It is still commonly found in decora- 
tions, but no figures are presently available concerning the 
amount used. The foliage of mountain laure] can also be 
used to dye cloth. Nicholson and Clovis (1967) found that 
the boiled leaves of this species produced a yellow tan color 
on wool if a chrome mordant was used. 

In the past, the wood of the mountain laurel was occa- 
sionally used to make small items. Peter Kalm in his journal 
(Benson, 1937) mentions that the strong wood is carved 
into weaver's shuttles, pulleys, spoons and trowels. Browne 
(1857) considered the wood to be similar to that of the 
European Box and that it was sometimes used for the 
handles of light tools, for screws, boxes, and that it was 
supposed to make good clarinets. He also mentioned that 
the American Indians used the wood for making small 
dishes, spoons, and other domestic utensils, and according 

320 Rhodora [Vol. 76 

to Benson (1937), this was the source of the common name 
"spoonwood." Other uses include the making of rustic 
furniture and as a substitute for brier pipes (Britton, 
1913). Due to the small size of most mountain laurel 
shrubs, the wood is rarely used at the present time except 
locally for tool handles and novelties. 

Kalmia latifolia, as a common understory shrub, is im- 
portant in preventing water runoff and soil erosion. Studies 
by Johnson and Kovner (1956) in the southern Appalachian 
Mountains show that the cutting of dense stands of laurel 
and rhododendron in hardwood stands decreased the evapo- 
transpiration rate and correspondingly increased the annual 
streamflow on the average of 2 inches per year. 

Extracts from the leaves of most species of Kalmia were 
at one time considered important as medicine, Barton 
(1802, 1804) mentions that a saturated tincture prepared 
from the leaves of Kalmia latifolia in a proof spirit could 
be used for certain fevers. According to Rafinesque (1830) 
preparations of this species were used as a narcotic, errhine, 
antisyphilitic and antiherpetic, while Griffith (1847) con- 
sidered it useful for fever, itching and in cases of hemor- 
rhaging and dysentery. Wood and Bache (1851) considered 
it useful mainly for cutaneous affections when applied as 
an ointment or decoction. During the latter part of the 19th 
century the leaves of K. latifolia were still commonly found 
in drug stores. According to Paschkis (1881), they were 
dried, rolled into a pipe-form and used principally as a 
remedy for diarrhea. The leaves of K. hirsuta were used 
in the south as a cure for itching and for mange in dogs 
(Porcher, 1869), while Holmes (1884) reports that K. 
angustifolia was commonly used by the Cree Indians of the 
Hudson Bay region for bowel complaints and as a tonic. 


Numerous cultivars of Kalmia latifolia have been named; 
no large scale nursery production of any of these forms 
exists, however, because they have been considered difficult 

1974] Genus Kalmia — Ebinger 321 

to root, hard to graft, and are untrue from seed. As a 
result, one common source of this species for ornamental 
purposes has been native woodland stands. Recent studies, 
however, indicate that it is possible to root cuttings of this 
species and that selected types do breed true. 

According to Jaynes (1971d) most Kalmia species can be 
started from cuttings. He found that greenwood cuttings 
of Kalmia angustifolia, K. hirsuta, K. polifolia, and K. 
microphylla root readily under mist or a plastic tent. By 
contrast, K. cuneata cuttings were difficult to root, while 
K. latifolia was highly variable in its rooting ability. In this 
latter species the ability to root is related to the age of the 
stock plant, since 89% of the cuttings from 1-year-old 
plants rooted, while only 21% of those from 4 or more 
year-old plants rooted. Jaynes also reported that most of 
the sterile hybrids can be propagated readily from cuttings, 
even when K. latifolia is one of the parents. Jaynes (1972) 
found that clones of K. latifolia vary in their ability to root 
from cuttings and that cuttings of cuttings or young grafts 
root more readily than cuttings from the original stock 

Few attempts have been made to graft any of the species 
of Kalmia. Trumpy (1893) had some success using side 
grafts with current year’s wood of Kalmia latifolia, while 
Jaynes (1971d, 1972) grafted current year’s growth of 
K. latifolia onto stock of the previous year using cleft or 
side-veneer grafts. The grafts were made in mid-June, 
kept in plastic tents for 6 weeks, hardened for 2 weeks, and 
overwintered in a lath house. The first report on the suc- 
cess of layering K. angustifolia was probably by Curtis 

The propagation of Kalmia from seeds has been exten- 
sively studied. Nichols (1934) found that Kalmia polifolia 
seeds germinated only after stratification, while K. angusti- 
folia, and K. latifolia seeds had a germination rate 5 times 
higher after stratification for 12 weeks. In contrast, Jaynes 
(1971c) reported that of these 3 species, only K. latifolia 
required stratification, the others germinating without any 

322 Rhodora [Vol. 76 

pregermination treatment. Jaynes also found that K. cu- 
neata, as well as K. latifolia, required stratification, but that 
soaking the seeds in gibberellin for 24 hours created the 
same response as stratification. The most difficult species 
to propagate appears to be K. hirsuta, which germinated 
best after a heat-humidity treatment (heating the seeds in 
a moist chamber at 80°C for 30-60 minutes). Jaynes 
(1968c) suggested that this response to high temperature 
may have adaptive value and could have evolved as a result 
of recurring ground fires. He also suggests that K. ericoides, 
a closely related species that is native to sandy pine areas 
of Cuba, may require a similar pregermination treatment. 
Kalmia microphylla is similar to K. polifolia in that it needs 
no pregermination treatment. The studies of Flemer (1949) 
and Jaynes (1971c) indicate that all species of Kalmia 
germinate best at 22°C under light in a mixture of sand, 
ground sphagnum, and peat moss that has a pH of 4.2. 

A mycorrhizal association, without which growth will 
not occur, has been demonstrated for Kalmia latifolia, and 
it is probably necessary for all species of the genus. Flemer 
(1949) found that mountain laurel seedlings grew very 
poorly in sterile culture without the associated endophyte, 
but when the seedlings were combined with the fungal 
associate they rapidly outgrew the controls. Microscopic 
examination revealed hyphae sheathing the younger por- 
tions of the roots and frequent hyphal connections with 
mycorrhizal coils in the cortical cells. No attempt was made 
to identify the fungus involved. 


Many species of Kalmia are sympatric, and most have 
similar habitat requirements. Even so, no natural hybrids 
have been reported, and recent studies suggest that such 
hybrids are unlikely to occur. Jaynes (1968a, 1971d) made 
all possible crosses, including reciprocals, between 6 species 
of the genus (omitting only Kalmia ericoides) and found 
that the barriers to gene flow are well developed. Most of 

1974] Genus Kalmia — Ebinger 323 

the interspecific crosses were difficult to complete since no 
seed was set, the resulting seed failed to germinate, or the 
seed produced albino or yellow-green seedlings. Stylar inhi- 
bition of pollen tube growth appeared to be the most com- 
mon reason for the lack of seed set. Only 9 combinations 
yielded viable seedlings and all were sterile. By contrast, 
intra-specific crosses between plants differing in flower color 
and growth habit within and between geographie sources 
showed no evidence of genetic barriers. 

Little is known about the inheritance of traits in the 
genus; the only detailed study involves flower color in 
Kalmia angustifolia. In this species, Jaynes (1971b) found 
that the presence of anthocyanin pigment is determined by 
a single dominant gene, with the homozygous recessive 
lacking red pigment in the leaves and stem and having 
white flowers. This trait is carried in the population as a 
heterozygote, but is rarely expressed. These white forms 
are at a selective disadvantage in the wild as the seedlings 
are weak, especially when small. 

Inbreeding depression and self incompatibility have been 
observed in some species of Kalmia. Jaynes (1968b) found 
that self pollinated Kalmia latifolia and K. angustifolia set 
85-90% less seed than cross pollinated plants, while K. poli- 
folia, in contrast, showed no self incompatibility. The seed- 
lings of all three species, however, showed inbreeding de- 
pression. As measured by height growth, the vigor of the 
outcrossed seedlings was generally twice that of the selfs. 
Jaynes (1971a) also noticed inbreeding depression when 
sibling red-budded mountain laurel was crossed. It is possi- 
ble that compact seedlings of Kalmia species of potential 
ornamental value can be obtained by selfing. 

Few cytological studies of the genus have been under- 
taken. Hagerup (1928) in an extensive cytological study 
of the bicornes reported a chromosome number of n = 12 
for Kalmia latifolia and n = 24 for K. glauca (K. poli- 
folia). He found that the chromosomes of K. polifolia were 
about half as large as those of K. latifolia, so that the 
amount of chromatin in the two species seemed to be about 

324 Rhodora [Vol. 76 

the same. Later, Callan (1941) reported a chromosome 
number of 2n — 44 for K. polifolia, which in light of recent 
studies is an error. Jaynes (1969) studied cytologically all 
of the taxa except for the Cuban species. He found that 
with the exception of K. polifolia (n — 24), all had a chro- 
mosome number of » — 12. No differences were found in 
the size of the chromosomes, and their smallness prevented 
a karyotype analysis. 


The most distinctive feature of the genus is its pollen- 
discharge mechanism. Near the middle of the corolla are 
10 sacs that form small lobes on each keel of the flower bud. 
Just before anthesis the anthers are pushed upward into 
these sacs by the elongating filaments. As the corolla opens 
the anthers are held in the sacs and are carried down and 
outward, the elastic filaments being bent backward under 
tension. When the flower is disturbed by a large insect, one 
or more of the anthers are released from their sacs. The 
filament, which is under tension, snaps the anther upward, 
showering the insect with pollen. This mechanism was first 
described by Sprengel (1793) and later by Hasskar] (1863) 
for Kalmia latifolia, while Rothrock (1867) observed the 
same mechanism in K. angustifolia, Studies by Penhallow 
(1882) showed that the pollen of K. latifolia could be 
thrown up to 14 cm. from the flower while in K. angusti- 
folia it was thrown up to 8 cm. away. These authors sug- 
gested that this was a method of insuring self-pollination 
since the pollen was thrown toward the stigma of the flower. 
Beal (1867) was the first to describe the way in which the 
flowers were cross-pollinated by insects. He observed that 
while a bumble bee was searching around the base of the 
flower, the proboscis liberated the stamens which projected 
their pollen onto the underside of the insect’s body. This 
pollen was then rubbed onto the stigma of other flowers 
visited. This process was also discussed by Gray (1876, 

1974] Genus Kalmia — Ebinger 325 

When an anther is released, the pollen tetrads are dis- 
charged as a fine powder in some species, and in others as 
a sticky net caused by the presence of viscid threads that 
hold the tetrads together. According to Matthews and Knox 
(1926) these extremely fine, non-cellular threads are prob- 
ably derived from small quantities of microspore mother 
cell protoplasm which are excluded from the pollen tetrads. 
They also suggest that this feature is an adaptation that 
facilitates pollination in relatively large, upright flowers 
like those of the genus Rhododendron. These threads were 
found in Kalmia latifolia by Ikuse (1954) and have been 
observed by the present author in K. hirsuta and K. eri- 
coides. Their presence or absence was used by Copeland 
(1943) as a major characteristic in separating the tribes of 
the subfamily Rhododendroideae, with the tribe Phyllodo- 
ceae being the only one that consistently lacked these 
threads. According to Wood (1961) it is now apparent that 
viscid threads do occur in some species of Kalmia, Phyllo- 
doce and Loiseleuria, all members of the Phyllodoceae. 
Because of this, the usefulness of this characteristic to 
distinguish tribes is doubtful. 

Only a few species of insects have been observed polli- 
nating Kalmia species. Beal (1867) listed bumble bees and 
other Hymenoptera, but mentions no species. The only de- 
tailed account was by Lovell and Lovell (1934) who col- 
lected and identified the pollinators found on a population 
of Kalmia angustifolia near Waldoboro, Maine. Their ob- 
servations show that K. angustifolia is effectively cross- 
pollinated by bees and that a total of 14 species of insects 
is capable of springing the stamens. Of these, Bombus 
ternarius and Andrena vicina were the most common visi- 
tors to the flowers. During the present study, populations 
of K. angustifolia and K. latifolia were examined in south- 
ern Connecticut, and all insects found to release the stamens 
were collected and identified. These studies were made for 
2 hour periods, at various times each day, from June 15 to 
June 25, 1970. In most instances the insect did not release 
the stamen by just landing on the flower. Usually the sta- 

326 Rhodora [Vol. 76 

mens were sprung by the proboscis while searching for 
nectar, but sometimes the legs set the stamens free. No 
honey bees were found on the flowers, and none were re- 
ported by Lovell and Lovell (1934). 

Recently bees have been used by Jaynes (1971a) to polli- 
nate caged plants of Kalmia latifolia and K. angustifolia for 
the large scale production of seed. In these experiments 
selected plants were caged before the flower buds opened. 
Honey bees (Apis spp.) or bumble bees (Bumbus spp.) 
were introduced into the cages when the flowers started to 
open. The plants that were in cages with bumble bees set 
large quantities of seeds (58 to 95% of the flowers forming 
capsules), while plants in cages with honey bees set fewer 
seeds (2 to 31% of the flowers forming capsules). Plants 
that were in cages with no bees rarely set seed (less than 
1% of the flowers forming capsules). 


Catesby (1743) was one of the first to report on the 
poisonous properties of members of this taxon. He found 
that when cattle and sheep were deprived of better food 
during severe winters, many died from eating the leaves of 
Kalmia latifolia (as Chamaedaphne). An extensive report 
by Kalm (Benson, 1937) mentions that young sheep die 
immediately or become very sick after eating leaves of 
Kalmia latifolia, but calves “were cured by giving them 
gunpowder and other medicines.” According to Kalm, 
larger domestic animals are also affected by the leaves. 

Most of the literature on livestock poisoning by Kalmia 
latifolia and K. angustifolia has been reviewed by Crawford 
(1908), Muenscher (1957) and Kingsbury (1964). Some 
experimental work has been done to determine the symp- 
toms, dosage, and treatment of the poisoning by these spe- 
cies. One of the first studies was made by Wood (1883), 
who fed a decoction of K. angustifolia leaves to a sheep. 
The symptoms were similar to those observed by Pritchard 
(1956) who studied K. angustifolia poisoning in a flock of 

1974] Genus Kalmia — Ebinger 327 

Minnesota sheep. Experiments with K. latifolia by Cary 
and Matthews (1903), Crawford (1908) and Marsh and 
Clawson (1930) showed that all livestock tested were poi- 
soned by this species. The western species, K. microphylla, 
is also poisonous but, because of its habitat, is seldom 
encountered by livestock. Fleming (1920) fed this species 
to some calves and sheep, most of which became sick, while 
Clawson (1933) found that the symptoms exhibited by 
sheep and goats were similar to those observed in animals 
fed K. latifolia and K. angustifolia. No information is 
available concerning the poisonous properties of the other 
members of the genus. 

Catesby (1743), Kalm (Benson, 1937), Bigelow (1818), 
and other early authors mentioned that deer can eat the 
leaves of mountain laurel without suffering ill effects. 
Forbes and Bechdel (1930, 1931) indicate that laurel is 
eaten in considerable quantity by deer, especially in time of 
food shortage. When restricted to a laurel diet, however, 
deer become thin and weak and usually develop a mild con- 
dition of rickets. Death results when the animals are fed 
in excess of 1.29% of their live weight of laurel leaves. 

All species of Kalmia are probably poisonous to humans 
(Hardin and Arena, 1969), although no deaths have been 
attributed directly to this genus. Bigelow (1818) and Grif- 
fith (1847) have reported instances of poisoning in humans. 
In these cases, a decoction made from the leaves of Kalmia 
latifolia produced toxic symptoms. The tough leaves and 
their bitter taste make it unlikely that anyone would eat 
them. Also, the quantity required to produce illness is 
greater than what a person would probably consume. Ac- 
cording to Barton (1802) the Delaware Indians used the 
leaves of K. latifolia for suicide. 

It has been suggested by various authors that the honey 
produced from Kalmia flowers is poisonous, but no definite 
proof is available. Barton (1802) reported that the honey 
from Kalmia angustifolia, K. latifolia, and K. hirsuta is 
poisonous to humans and listed a few instances of suspected 
poisoning by honey of these species. Miller (1895) and 

328 Rhodora [Vol. 76 

Pellett (1920), in contrast, doubted that mountain laurel 
was responsible for poisonous honey. They felt that due to 
the abundance of this species it would seem that the cases 
of poisoning would be reported much more frequently if 
there was good reason to suspect the honey from this source. 
Some doubt is also expressed by Lovell (1926, 1951) since 
bees are rarely found on mountain laurel. Howes (1949), 
as well as Hardin and Arena (1969), indicate that Kalmia 
latifolia honey is an extremely distressing narcotic, varying 
in its effect in proportion to the quantity eaten. In none of 
the reports is the source of the poisonous honey known for 
certain, the genus Kalmia being suspected because other 
parts of these plants are known to be poisonous. 

Andromedotoxin, the poisonous compound found in the 
genus Kalmia, is known to occur also in other genera of the 
Ericaceae. Numerous attempts have been made to isolate 
and determine the chemical structure of this compound. 
Lasché (1889) reported the occurrence of andromedotoxin 
in the leaves and twigs of Kalmia angustifolia and also in 
the fruits of K. latifolia, while Matusow (1897) found what 
he considered andromedotoxin in the roots of K. latifolia. 
More recently Jacobs and Lloyd (1939) and Waud (1940) 
isolated andromedotoxin from K. angustifolia. The chemical 
structure of andromedotoxin has been studied by Wood, 
Stromberg, Keresztesy and Horning (1954) and more re- 
cently by Tallent, Riethof and Horning (1957). The latter 
authors renamed the substance acetylandromedol and de- 
veloped a procedure for detecting its presence in plant 
extracts through the use of paper electrophoresis. Though 
the chemical structure of andromedotoxin is still in doubt, 
its empirical formula is probably C..H,,0.. 


Only a few fossil species of Kalmia have been named. 
Most are described from leaf impressions, and except for 
size, shape, and probable coriaceous texture, there is little 
reason to consider them members of the genus Kalmia, or 

1974] Genus Kalmia — Ebinger 329 

members of the family Ericaceae. One living species is also 
known from fossil remains. 

Kalmia brittoniana Hollick (1892). 

This fossil was first described from clay beds of the 
Raritan formation, near Kerischerville, Staten Island, New 
York and is Upper Cretaceous in age (Museum, Staten 
Island Association of Arts and Science). The material il- 
lustrated consists of parts of three oblong, entire, probably 
coriaceous leaves, 0.7 X 2.7 cm., with prominent midribs 
and no secondary veins. Hollick (1906) mentions that the 
absence of secondary nervation, indicating a leaf of thick, 
coriaceous texture, was what largely influenced his refer- 
ring the leaf to the genus Kalmia. This species has also 
been reported from the Upper Cretaceous deposits of North 
Carolina (Berry, 1907, 1920) and Alabama (Berry, 1913, 
1919). When he described Kalmia brittoniana from the 
Tuscaloosa formation of Alabama, Berry (1919) suggested 
that there is no reason for its placement in the genus 
Kalmia, or even in the Ericaceae. He further sugested that 
the fossil probably represents some Upper Cretaceous spe- 
cies of the Proteaceae or the Myrtaceae. 

Kalmia ? elongata Ashlee (1932). 

This species was found in the Latah formation, Elk River, 
Idaho, and is Upper Miocene in age (University of Idaho 
collection No. 303). The specimen consists of two narrowly 
elliptical, subcoriaceous leaves, 0.9 X 1.7 cm., with entire 
margins and a petiole 2-4 mm. long. The leaves are super- 
ficially similar to Kalmia microphylla, but since the midrib 
is grooved and the secondary and tertiary veins are present, 
this fossil must be considered a doubtful member of the 

Kalmia elliptica Brown (1962). 

This species was found in the Fort Union formation (Ilo 
Post Office, Wyoming) and the Denver formation (Golden, 
Colorado) and is Paleocene in age. The specimen illustrated 
by Brown (1962) consists of a single elliptic leaf with 
entire margins and secondary and tertiary veins that ir- 

330 Rhodora [Vol. 76 

regularly fork and loop near the margins. The specimen is 
similar in outline to a typical leaf of Kalmia latifolia but 
is about twice as large (7 X 17 cm.). The presence of the 
well developed and large number of secondary and tertiary 
veins, as well as their angle to the midrib, is not typical of 
any present day species of Kalmia. It must therefore be 
considered a doubtful member of the genus. 

Kalmia saxonica Litke (1968). 

This species was found at Brandis, northwest Saxony, 
Germany, and is Lower Miocene in age. It consists of pieces 
of cuticle with the upper epidermis occasionally attached. 
'These fragments indicate that the leaf was entire margined, 
1-1.3 em. wide, and had an obvious midvein. The cells are 
polygonal with wavy margins, the stomata are round, 18u 
wide, surrounded by a double cuticular ring and occasionally 
by radiating cuticular striations. The basal portion of glan- 
dular hairs are also present. The structure and arrange- 
ment of the cells is similar to that found in Kalmia latifolia, 
and it is possible that this fossil represents a member of 
this genus or a closely related genus of the Ericaceae. 
Similar cuticle remains were placed in the form genus 
Kalmiophyllum by Kráusel and Weyland (1959) when they 
described the species Kalmiophyllum marcodurense. 

Kalmia polifolia, the Bog Laurel, has also been found in 
fossi] form. It was first reported by Berry and Johnston 
(1922, as K. glauca Ait.) from interglacial deposits at 
Point Grey near Vancouver, British Columbia. The fossil 
specimen illustrated is of the same shape and size as those 
of living specimens of K. polifolia. Since the other fossil 
plants found clearly indicate a bog habitat, it is very pos- 
sible that this fossil is a species of Kalmia. Kalmia polifolia 
has also been reported from Pleistocene lake deposits of the 
upper Connecticut River Valley in northern New Hamp- 
shire (Hollick, 1931). These fossils are post-glacial in age 
and according to Hollick are representative of the flora that 
migrated northward in the wake of the retreating Wiscon- 
sin ice sheet. The other fossil species found in the same 
deposits indicate a habitat and climate similar to that pres- 

1974] Genus Kalmia — Ebinger 331 

ently prevailing in the area. The plates in this article show 
excellent examples of leaves very similar in shape and size 
to those of living K. polifolia. As mentioned by Hollick, 
however, positive identification would “merely represent a 
matter of personal opinion" since the leaves of K. polifolia, 
Andromeda polifolia, and Ledum groenlandicum appear al- 
most identical in outline. 


KALMIA Linnaeus, Sp. Pl. 1: 391. 1753. 
Kalmiella Small, Fl. Southeast. U. S. 886. 1908. 
Chamaedaphne Catesby ex Kuntze, Rev. Gen. Pl. 2: 388. 
1891, non Moench. 

Evergreen or rarely deciduous shrubs or small trees with 
pubescent to occasionally glabrous, terete or 2-edged twigs. 
Lateral buds very small, flattened, with 2 exposed valvate 
to overlapping scales. Leaves alternate, opposite, or in 
whorls of 3, exstipulate, usually coriaceous, petioled to sub- 
sessile, margins entire. Flowers solitary in the axils of 
leaves, or in terminal or axillary corymbiform racemes, or 
in terminal compound corymbs. Pedicels slender, terete, 
usually pubescent and with stalked glandular hairs, sub- 
tended by a pair of bractlets in the axil of a small leaf-like 
bract. Flowers 5-merous, actinomorphic, with hypogynous 
insertion. Calyx synsepalous, usually green and persistent 
in fruit; lobes 5, longer than the tube. Corolla sympetalous, 
white to pink, red or purple, with a short cylindrical tube 
extending into a rotate limb that is shallowly 5-lobed and 
commonly 10-keeled, the limb with 10 saccate depressions 
in which the anthers are held under tension. Stamens 10, 
shorter than the corolla; filaments usually filiform, glabrous 
or pubescent, dorsifixed; anthers 2-celled, unappendaged, 
short, ovoid, opening by large terminal slits; pollen dis- 
charged in tetrads, with or without viscid strands. Ovary 
superior, 5-celled, placentation axile, placentae massive with 
numerous ovules; style slender, terete, nearly straight; 
stigma unexpanded, slightly grooved. Fruit a subglobose 

332 Rhodora [Vol. 76 

to globose, 5-valved, septicidal capsule usually subtended by 
the persistent calyx. Seeds numerous, small and light- 
weight, testa tight or loose and extending well past the 
ends of the remainder of the seed. A North American 
genus extending from Alaska south in the mountains to 
California and Utah, eastward through Canada to the At- 
lantic Ocean and southward through eastern United States 
to Florida and Cuba. 

Lectotype Species: Kalmia latifolia L. (cf. Britton 1908). 


A. Leaves opposite; inflorescence a terminal corymbiform 

B. Midrib of leaves lacking stalked glands; seeds less 
than 1.5 mm long . .. ..... 1. K. microphylla. 

B. Midrib of leaves with stalked glands on both sur- 
faces; seeds more than 1.5 mm long. . 2. K. polifolia. 
A. Leaves alternate or in whorls; inflorescence various. 

C. Inflorescence a terminal compound corymb; leaves 

usually more than 2 em broad. ..... 3. K. latifolia. 

C. Inflorescence a simple corymbiform raceme or the 

flowers solitary in the axils of the leaves; leaves 
usually less than 2 cm broad. 

D. Leaves more than 1.5 em long; inflorescence a 

corymbiform raceme from the axils of the leaves. 

E. Leaves in whorls of 3, evergreen. .......... 
IDEM ... 4. K. angustifolia. 

E. Leaves alternate, deciduous. .. 5. K. cuneata. 

D. Leaves less than 1.5 cm long; flowers mostly soli- 
tary in the axils of the leaves. 

F. Leaves broad, flat, the margins only slightly 
revolute; flowers scattered along the stem in 
the leaf axils. .............. 6. K. hirsuta. 

F. Leaves narrow, strongly revolute; flowers 
clustered toward the ends of the twig giving 
the appearance of a terminal corymbiform 
raceme. ........... ls 7. K. ericoides. 

1974] Genus Kalmia — Ebinger 333 

1. Kalmia microphylla (Hooker) Heller, Bull. Torrey Bot. 
Club 25: 581. 1898. 

Low growing alpine plant to sparingly branched shrub 
to 6 dm tall; branchlets terete to slightly 2-edged below the 
leaves, glabrous to lightly pubescent; leaves opposite, sub- 
sessile, the broad petiole to 2 mm long, the blades coriaceous, 
ovate to oval to narrowly lanceolate, to 4 em long, 0.4-1.1 cm 
broad, whitened beneath with small hairs, a band of white 
hairs usually present on the upper surface on either side 
of the midrib, midrib lacking stipitate glands, margins flat 
to slightly revolute; inflorescence a few-flowered, terminal, 
racemose cluster, each flower on a glabrous pedicel 1-3 cm 
long, each pedicel subtended by a pair of bractlets in the 
axis of a leaf-like bract; calyx pink to light green, 5-10 mm 
broad, the lobes obtuse, margins ciliate; corolla rose-purple 
(rarely white), 0.8-2.0 cm broad; anthers purple, 0.6-1.6 
mm long; fruit subglobose, 4-7 mm broad, 3.5-6 mm long; 
seeds 0.8-1.4 mm long, the testa extending past the ends of 
the seed. Chromosome number n — 12. 

Kalmia microphylla has rarely been considered specifically 
distinct from K. polifolia, Hall (1912), when he transferred 
the taxon to a variety of K. polifolia mentioned that “No 
characters upon which the variety might be raised to spe- 
cific rank have as yet been pointed out, except the variable 
one of size in plant and leaf." Most present floristic works 
maintain this taxon as a variety or subspecies of K. poli- 
folia, though it has been considered a separate species by 
Small (1914), Rydberg (1954), Hultén (1948) and Abrams 
(1951). Hultén (1948) mentioned that it was with some 
hesitation that he accepted K. microphylla as being specifi- 
cally different from K. polifolia, but that in Alaska, there 
does not seem to be any intermediate types. However, in 
1968 he considered K. microphylla a subspecies of K. poli- 

When these taxa are considered distinct the characters 
of plant and leaf size are the most commonly used for 
separation. Kalmia polifolia is a large shrub, 3 or more dm 

384 Rhodora [Vol. 76 

tall, the leaves are 2-4 cm long, 214 to 3 times longer 
than broad, and the margins are usually strongly revolute. 
Kalmia microphylla, in contrast, is a much smaller plant, 
usually less than 1.5 dm tall, the leaves are usually less than 
1.5 em long, less than 215 times as long as broad, and the 
margins are little if at all revolute. In most instances, these 
characters are sufficient to separate the two taxa; however, 
variation and overlapping of characters do occur. Other 
characters that have previously been overlooked are much 
more reliable in separating the two species. In K. polifolia 
clavate glands are present on the upper and lower surface 
of the leaf midrib, and the seeds are 1.5 to 2.2 mm long, 
while in K. microphylla these glands are absent, and the 
seeds are usually less than 1.3 mm long. 

Kalmia microphylla is the only species of the genus found 
west of the Rocky Mountains. It extends from central 
California north to Alaska, and east through the Yukon 
and Northwest Territories to the extreme northwest corner 
of Manitoba. It is primarily a low-growing alpine shrub 
rarely exceeding 1 dm in height, but in bogs at lower eleva- 
tions it may reach a height of 6 dm. The species consists 
of two varieties, one restricted to alpine and tundra areas, 
and one found in coastal lowland regions. 


1. Leaves oval to broadly elliptic, usually less than 2 cm 
long and less than 214 times as long as broad; calyx 
less than 7 mm broad. .......... la. var. microphylla. 

1. Leaves lanceolate, usually 2.5 cm long or longer, 21% to 
4 times as long as broad; calyx over 7 mm broad. .... 

gaa new wea nEy ecaey eenerhu names 1b. var. occidentalis. 

1a. Kalmia microphylla (Hooker) Heller var. microphylla. 

Kalmia glauca Ait. y microphylla Hook. Fl. Bor.-Amer. 2: 
41. 1834. TYPE: swamps in the Rocky Mountains, Drum- 
mond (K). Kalmia polifolia var. microphylla Rehder in 
Bailey, Cycl. Am. Hort. 2: 854. 1900. (without basionym). 
Kalmia polifolia var. microphylla (Hooker) Hall, Univ. 

1974] Genus Kalmia — Ebinger 335 

California Publ. Bot. 4: 201. 1912. Kalmia polifolia ssp. 
microphylla (Hooker) Calder & Taylor, Canad. Jour. Bot. 
43: 1398. 1965. 

Sparingly branched alpine shrub usually less than 1 dm 
tall (to 3 dm tall at lower elevations) ; nodes usually less 
than 1 em apart; leaves ovate to oval to broadly elliptic, to 
1.5 (rarely 2.5) cm long, less than 214 times as long as 
broad; calyx pink to light green, 5-7 mm broad; corolla 
rose-purple 0.8-1.2 (rarely 1.5) cm broad; fruit 4-5 mm 
broad, 3.5-4.5 mm long; seeds 0.8-1.3 mm long. 

Distribution: Mountainous regions of western North 
America from California, Nevada, Utah and Colorado north 
through the Rocky Mountains to the Yukon and Northwest 

Common name: Alpine Laurel, Small-leaved Kalmia. 


UNITED STATES: Alaska: Skagway, Anderson 1608 (1sc), 
Eastwood (GH). California: EL DORADO CO.: Echo Lake, Beatle & 
Beatle 3959 (ARIZ, DUKE, NY). FRESNO CO.: Marie Lake, Kruckeberg 
3487 (RM, WS). HUMBOLDT CO.: Trinity Summit, Davy & Blasdale 
5823 (Us). INYO CO.: Pine Creek, Duran 3338 (GH, MICH, US). 
MADERA Co.: Shadow Creek, Raven (NY). MARIPOSA CO.: Crescent 
Lake, Congdon (MIN, WS). MODOC CO.: Dismal Swamp, Weatherby 
1598 (NY, RM). MONO CO.: Treble Lake, Munz 19998 (NY). NEVADA 
co.: Frog Lake, Sonne 438 (MU, PH, US). PLACER CO.: Squaw Valley, 
Eggleston 21607 (GH, NY, US, WS). PLUMAS C0.: Long Lake, Hall 
9325 (GH, RM, US). SHASTA C0.: Lassen Nat Park, Ferris & Lorraine 
10459 (GH, RM, US). SISKIYOU CO.: Mt. Shasta, Cooke 11296 (GH, 
osc), 17654 (WS, WIU). TULARE CO.: Sky Blue Lake, Howell 25555 
(US). TUOLUMNE CO.: Unicorn Creek, Mason 698 (GH). Colorado: 
BOULDER C0.: Brainard Lake, Green 279 (PENN). CLEAR CREEK CO.: 
Loveland Pass, Robbins 732 (ARIZ). GILPIN C0.: Forest Lake, Roberts 
(CM). JACKSON CO.: Cameron Pass, Beaman & Erbisch 1362 (MSC, 
RM). JEFFERSON CO.: Geneva Creek, Livingston 298 (DUKE). LARIMER 
co.: Estes Park, Cooper 116 (RM), 157 (ARIZ, MIN, NMC, RM). 
ROUTT CO.: Routt Natl Forest, Weber 6991 (ws). Idaho: BLAINE 
co.: Sawtooth Peaks, Macbride & Payson 3707 (GH, RM, US). 
BOISE CO.: Middle Spangle Lake, Hitchcock & Muhlick 10130 (GH, 
NY, RM). BONNER CO.: Priest Lake, MacDougal 292 (NY). BOUNDARY 
co.: Mt. Roothaan, Daubenmire 44289 (WTU). CLEARWATER CO.: 
Lolo Trail Rd., Joslyn 953 (1D), 954 (ID). FREMONT CO.: Henry Lake, 

336 Rhodora [Vol. 76 

Payson & Payson 2047 (cH, NY, RM). IDAHO CO.: Big Fog Mt. 
Baker 14762 (1D). SHOSHONE co.: Little Lost Lake, Wilson 318 
(GH, IDS, WS). VALLEY CO.: Boulder Lake, Boone 177 (1D). Montana: 
BEAVERHEAD CO.: Odell Lake, Hitchcock & Muhlick 14915 (NY, WS, 
WTU). CARBON CO.: Beartooth Mts., Cronquist 8068 (GH, NY, ws, 
Standley 18513 (US). GALLATIN CO.: Spanish Peaks, Vogel (cu, 
MONT, US). GLACIER CO.: Hidden Lake, Hitchcock 1905 (MONT, RM). 
GRANITE CO.: Helena, Kelsey (MU, NY). LAKE CO.: McDonald Lake, 
Hitchcock 18343 (RM, ws, WTU). MADISON C0.: Old Hollowtop, 
Rydberg & Bessy 4654 (MONT). MISSOULA CO.: Lace Lake, Rossbach 
& Rossbach 184 (WTU). PARK CO.: Lady of the Lake, Witt 1233 
(MINN, NY, WS, WTU). POWELL CO.: Upper Tineup Creek, Trask 117 
(MONT). RAVALLI CO.: St. Mary's Peak, Hitchcock & Muhlick 15332 
(NY, WS, WTU). STILLWATER CO.: Stillwater, Hawkins 35502 (MONT). 
SWEET GRASS CO.: Rainbow Lake, Hitchcock 16501 (CAN, IDS, NY, WS, 
WTU). TETON C0.: Henry Mts., Williams (GH). Nevada: ELKO Co.: 
Lamoille Lake, Holmgren 1404 (BRY, NY). WASHOE Co.: Mt. Rose, 
Archer 6425 (ARIZ, GH, NY, RENO). Oregon: BENTON Co.: Three 
Sisters, Vechten 212 (osc). CLACKAMAS CO.: Clackamas Lake, Bart- 
lett & Grayson 791 (NY). CROOK COo.: Des Chutes Nat'l. Forest, 
Ferris & Duthie 488 (RM). DESCHUTES CO.: Sparks Lake, Bellinger 
(WILLU). DOUGLAS CO.: Umpqua, Ingram 1515 (ORE). JACKSON CO.: 
Mt. Pitt, Peck 5184 (WILLU). KLAMATH CO.: Crater Lake Applegate 
319 (US, wILLU), 345 (Us). LANE Co.: McKenzie Pass, Henderson 
16315 (ORE). LINN C0.: Hensley Meadows, Coville & Applegate 1152 
(US). MARION CO.: Russell Lake, Lee 44 (osc). UNION co.: Head 
1298 (osc). WALLOWA CO.: Mirror Lake, Peck 1777 (NY, WILLU). 
Utah: DUCHESNE CO.: Mt. Agassiz, Cottam & Biddulph 3703 (BRY). 
SUMIT CO.: Motte Peak, Payson & Payson 5139 (CHRB, GH, PH, RM, 
US, WS). UINTAH CO.: Leidy Peak, Maguire 17679 (RM). UTAH CO.: 
Grandaddy Lakes Basin, Stutz 224 (BRY). Washington: CHELAN CO.: 
Ice Lake, Morrill 332 ( MONT, ND, WTU). KITTITAS CO.: Mt. Stuart, 
Elmer 1103 (GH, K, MIN, US, WS). KLICKITAT CO.: Falcon Valley, 
Suksdorf (PH, ws, WTU). LEWIS CO.: Reflection Lake, Townsend 
(WS). OKANOGAN CO.: Slate Peak, Ownbey & Meyer 2302 (an, IDs, 
ISC, MIN, NY, OSC, US). PEND OREILLE CO.: Sullivan Lake Dist., 
Layser 866 (WS). PIERCE CO.: Mt. Rainier, Allen 96 (CAN, GH, K, 
MIN, MSC, NY, US, WS). SKAMANIA CO.: Chiquash Mts., Suksdorf 
(WS). WHATCOM CO.: Mt. Baker, Muenscher 8077 (ws). YAKIMA 
co.: Bird Creek Meadows, Zuberbuhler (ws). Wyoming: ALBANY 
co.: Brooklyn Lake, Nelson 9226 (GH, MIN, RM). BIG HORN CO.: 
Upper Crater Lake, Lofgren 13 (MIN, NY, RM). CARBON CO.: Silver 
Lake, Henry (CM). FREMONT CO.: Gannett Creek, Jozwik 386 (Gu, 
RM). JOHNSON CO.: Powder R. Pass, Stephens 40827 (VDB). LARIMER 

1974] Genus Kalmia — Ebinger 337 

co.: North Park Range, Goodding 1839 (PENN). PARK CO.: Island 
Lake, Truman 54298 (WTU). SHERIDAN CO.: Dome Lake, Pammel & 
Stanton 62 (MIN). TETON CO.: Taggart Lake, Williams 1119 (ID, 
Nelson & Nelson 6170 (CM, ISC, MIN, GH, NY, RM, US). 

CANADA: Alberta: Banff Natl. Park: Bow Pass, Saunders 11 
(rRT); Lake Louise, Brown 578 (GH, PH, US), McCalla 2162 (ALTA, 
NY, US); Waterfowl Lake, Walker 2443 (PENN). Jasper Nat. Park: 
Athabasca Falls, Moss 2752C (ALTA, GH); Maligne Lake, Scamman 
2664 (GH); Amethyst Lake, MacFadden 3246 (No). Waterton Lakes 
Nat! Park: Summit Lake, Moss 975 (GH, NY, US); Upper Twin 
Lake, Blais & Nagy 1793 (CAN). Fairfax Lake, Damais & Anderson 
2335 (ALTA); Mercoal, Woollven 6 (ALTA); Viewpoint, Damais & 
Kempinsky 5683 (ALTA); Virginia Hills, Moss 12469 (ALTA), 12505 
(ALTA, CAN). British Columbia: Ernest C. Manning Prov. Park: 
Timberline Valley, Hardy 18501 (usc, v); Three Brother Mts., 
Underhill 782 (v). Garibaldi Prov. Park: Castle Towers, Davidson 
(usc). Glacier Nat'l. Park: Bald Mt., Fogg, Jr. 1273 (PENN, UBC); 
Eagle Mt., Rosendahl 944 (MIN). Kokanee Glacier Prov. Park: 
Thompson 14447 (CAN, GH, MICH, NY, PH, US, WTU). Kootenay Natl. 
Park: Ross Lake, McCalla 9603 (uBC, v); Emerald Lake, Brown 
(TRT). Mt. Assiniboine Prov. Park: Magog Lake, Green 551 (PENN). 
Mt. Revelstroke Nat. Park: Calder & Savile 10861 (Ny, US), 
8884 (Nv, US). Tweedsmuir Prov. Park: Trew 17177 (v). Wells 
Gray Prov. Park: Battle Mt., Ahti & Ahti 6975 (v, wTU). Yoho 
Nat'l Park: Emerald Lake, Beer (cM); Selkirk, Shaw 41 (GH, 
MIN, NY, PENN, US); Yoho Lake, Pease 22371 (MASS). Vancouver 
Island: Mt. Arrowsmith, Taylor 164 (TRT); Mt. Albert Edward, 
Hett 352 (v); Moat Lake, Underhill 299 (wTU). Atlin, Szezawinski 
(UBC, V). Caribou Pass, Henry 608 (GH, ISC); Ft. Steele, Johnstone 
(UBC, v); Lillooet, Beamish & Vrugtman 610710 (vBC); Mt. Hazel, 
Slocomb (v); Penticton, Calder & Savile 10751 (NY, RM, UBC, V, WS) ; 
Prince George, Holland (v); Sikanni Chief R., Henry 743 (PH); 
Summit Lake Landing, Weber 2589 (GH, Ws); Skagit Valley, Macoun 
72406 (ND, NY); Tenquille Lake, Beamish & Vrugtman 61010 (1D, 
UBC, WTU). Manitoba: Churchill, Beckett M-45-5 (MIN), Ellis 1287 
(Us), Gillett 1832 (Isc, us), Gillis 3244 (msc), Voss 8999 (MICH), 
Walker 5090 (WIN); Tidal, Brown 178 (CAN, SASK). Northwest 
Territories: Aberdeen Lake, Macpherson 327 (CAN); Beverly Lake 
cabin, Kuyt 107 (CAN); Contwoyto Lake, Thieret 9415 (GH, US); 
Ennadai Lake, Brown 1320 (CAN): Gordon Lake, Henderson 19 
(CAN); Great Bear Lake, Porsild 17117 (cAN), Porsild & Porsild 
3429 (CAN), 3509 (CAN, US); Great Slave Lake, Pike (K); Mac- 
Millan Pass, Porsild & Breitung 11258 (CAN, GH, US) ; Maguse Lake, 
Gussow 33 (CAN); Nueltin Lake, Harper & Porsild 2211 (CAN, PH), 

338 Rhodora [Vol. 76 

2246 (CAN), 2271 (MICH, MIN, PH), 2554 (MICH, MIN, NY); Small 
Tree Lake, Maini & Swan 506 (RM, SASK); Thelon Game Sanctuary, 
Carroll 65 (CAN); Yellowknife Highway, Thieret & Reich 7742 
(CAN, MIN), 8121 (Us). Yukon: Granite Mt., Porsild & Breitung 
10576 (CAN, ISC, GH, US, WTU); Mayo Dist., Bostock 102 (CAN); 
McQueston Area, Campbell 493 (CAN); Mt. Sheldon, Porsild & Brei- 
tung 11094 (CAN, GH, US, WTU), 11485 (CAN, GH); pass between 
Teslin & Nisutlin R., Porsild & Breitung 11045 (CAN, Ny), 11148 
(CAN); St. Elias Mt., Pearson & Porsild 67-166A (CAN); Yukon- 
Mackenzie border, Calder & Kukkonen 27664 (Isc, NY); White Pass 
Summit, Bolton (us). 

Plant and leaf size in Kalmia microphylla var. micro- 
phylla is controlled to some extent by the environment. In 
the typical alpine form the leaves are extremely small, 
usually less than 1.2 em long. The type specimen, which 
represents this extreme, has leaves less than 1 cm long and 
is extremely short, less than 1 dm tall. At lower elevations 
the leaves are not uncommonly 2 em long and the plant 2 dm 
tall. Rarely taller plants are found with leaves to 2.6 cm 
long. These larger leaved forms are superficially similar to 
K. polifolia and probably account for the variation in tax- 
onomic treatment of K. microphylla in the past. 

Kalmia microphylla and K. polifolia probably never come 
into contact with each other. In the Northwest Territories, 
however, the northern part of both their ranges, it is possi- 
ble that the two taxa may occur together. The author has 
seen very few specimens of K. polifolia from the Northwest 
Territories, but this species is common in northern Alberta, 
Saskatchewan, and Manitoba. A few specimens of K. micro- 
phylla have been examined from the Northwest Territories, 
and this species extends all the way to the northwestern 
corner of Manitoba. No specimens, however, have been 
observed from Alberta or Saskatchewan. Sterile hybrids 
have been produced by crossing these two taxa (Jaynes, 
1968a), but no specimens from northern Canada have been 
found that are morphologically similar to these hybrids. 

The only other taxon with which Kalmia microphylla var. 
microphylla might be confused is K. microphylla var. occi- 
dentalis. Both taxa occur in western North America and 

1974] Genus Kalmia — Ebinger 339 

their ranges overlap. The two are easily separated, how- 
ever, since var. occidentalis has lanceolate leaves that are 
usually more than 2.5 cm long and 214-4 times as long as 
broad, while var. microphylla has oval to eliptical leaves 
that are smaller and less than 215 times as long as broad. 
Also, var. occidentalis has larger flowers, with the corolla 
more than 1.5 em broad and the calyx 7-10 mm wide. Vari- 
ety microphylla, by contrast, has smaller flowers, with the 
corolla less than 1.5 cm broad (usually less than 1.2 cm 
broad) and the calyx less than 7 mm wide. 

Although both varieties are found in marshes, bogs, and 
other open wet areas, it appears that they rarely come into 
contact with one another. Of the var. microphylla speci- 
mens examined, most were collected at relatively high 
elevations aecording to the information available on the 
herbarium labels. The elevation at which this taxon was 
collected varied from an average altitude of 8,500 feet above 
sea level in California (5,200 to 11,500), to about 10,000 in 
Colorado, Nevada, Utah and Wyoming (7,000 to 12,500), 
to about 6,500 in Oregon (5,800 to 7,700), and to about 
6,000 in Washington, Alberta and British Columbia (3,000 
to 7,300). The altitudes in parentheses indicate the mini- 
mum and maximum elevations recorded from the labels. 
The information on the labels of specimens of var. occi- 
dentalis indicates that this species is found at lower eleva- 
tions. Most of the specimens were from at or near sea level, 
but a few had been collected as high as 3,000 feet. 

Kalmia microphylla var. microphylla and K. microphylla 
var. occidentalis apparently hybridize when they come into 
contact. A few specimens from Vancouver Island and the 
Cascade Mountains of Washington appear to be intermedi- 
ate morphologically. These intermediate individuals are 
robust plants, 2-4 dm tall, similar in habit to the large- 
leaved var. microphylla. The leaves are 1.8-2.5 cm long, 
usually more ovate than lanceolate, and about twice as long 
as broad. The flowers are large, more typical of var. occi- 
dentalis. A few hybrids have been produced by crossing 
these varieties and they are highly fertile and set large 

340 Rhodora [Vol. 76 

quantities of seed (Jaynes, personal communication). Of the 

few suspected hybrids found, the pollen stainability varied 

from 10-90% normal pollen, indicating that some sterility 

occasionally results. 

1b. Kalmia microphylla var. occidentalis (Small) Ebinger, 
coinb. et stat. nov. 

Kalmia occidentalis Small, N. Am. Fl. 29: 53. 1914. 
TvPE: Washington: foothills of Mt. Rainier, 1885, Mrs. 
Bailey Willis (NY). Kalmia polifolia subsp. occidentalis 
(Small) Abrams, Ill. Fl. Pacific St. 3: 302. 1951. 

Sparingly branched shrub to 6 dm tall; leaves lanceolate, 
flat to slightly revolute margins, 2-4 cm long, some of the 
leaves always over 2 cm long and 215-4 times as long as 
broad; calyx light pink, 7-10 mm broad; corolla usually 
rose-purple (rarely white), 1.5-2.0 cm broad; fruit 5-7 mm 
broad, 4.5-6 mm long; seeds 1-1.4 mm long. 

Distribution: Along the western coast of North America 
from southern Alaska south to Oregon. 

Common name: Western Swamp Kalmia; Western Bog 



UNITED STATES: Alaska: Admiralty Isl., Hibber 236 (UNM); 
Annette Isl, Coville & Kearney 381 (vs); Douglas Isl, Thorne 
10280 (UWM); Gravina Isl., Anderson 5352 (1sc), 24599 (CAN, ISC); 
Kuiu Isl, Rigg 17 (wTU); Metlakatlan Isl, Emgstrom (WTU); 
Mitkof Isl, Brown (ALA); Pleasant Isl, Reiners (HNH); Prince of 
Wales Isl., Flett 1976 (vs); Revillagigedo Isl., McCabe 8590 (WTU); 
Woronkofski Isl., Klein 486 (ALA); Yakobi Isl., Anderson 1389 (US); 
Juneau, Anderson 6338 (ALA, BRY, CAN, GH, ISC, PH, RM, V) ; Menden- 
hall, Anderson 88 (CHRB); Petersburg, Scamman 515 (ALA, GH); 
Sitka, Anderson 160 (1sc, US); Wrangell, Walker & Walker 638 
(CM, GH, NY, RM, US, ws); Yes Bay, Gorman 21 (K, NO, NY, US). 
Oregon: CLATSOP CO.: Gearhart, Adair (osc); Knappa, Bouch 
(WTU); Delmoor, Cooke 10828 (osc). Washington: GRAY HARBOUR 
co.: Lake Quinault, Thompson 11429 (ALTA, GH, MONT, NY, PH, 
US, WS, WTU). ISLAND CO.: Whidbey Isl, Smith 480 (ws, WTU). 
KING CO.: Green Lake, Benson 1474 (WTU). MASON CO.: Lost Lake, 
Jones 6527 (PH, V, WTU). PACIFIC CO.: Ilwaco, Foster 865 (ORE, US). 
PIERCE C0.: Tacoma, Sloan (WTU). SAN JUAN CO.: Mt. Constitution, 
Zeller & Zeller 1193 (CM, GH, MIN, NY, US). SKAGIT CO.: Mt. 

1974] Genus Kalmia — Ebinger 341 

Higgins, Thompson 9639 (ND, US, WTU). SNOHOMISH CO.: Marys- 
ville, Grant (ARIZ, OAC, MU, RM, US). THURSTON CO.: Upper Valley 
of the Nisqually, Allen 4 (CAN, K, MIN, MSC, NY, US, WS). WHATCOM 
co.: Custer, Muencher & Muencher 6007 (PH, Us, WS). 

CANADA: British Columbia:  Aristazabal Isl, McCabe 3424 
(wTU); Dewdney Isl, Schofield 70-27 (UBC); Goose Isl, Martin & 
Guiquet (v); Hope Isl., Calder & Mackay 31240 (cM, MIN); Lulu Isl., 
Beamish & Vrugtman 60519 (Ny, osc, UBC); Graham Isl., Calder & 
Savile 21349 (osc, TRT, ws); Kunghit Isl., Foster & Bigg (UBC); 
Langara Isl., Beebe (v); Moresby Isl., Calder & Taylor 35771 (UBC); 
Prince Royal Isl, McCabe 3358 (wTU); Saltspring Isl, Ashlee 
(UBC, V); Vancouver Isl.: Comox, Anderson 483 (v); McCoy Lake, 
Carter 473 (us); Lost Lake, Macoun 618 (CAN, US); Port Hardy, 
Racey (UBC); Trout Lake, Davidson 6192 (usc); Alice Arm, 
Szczawinski (V); Alouette R., Garman (v); Garibaldi Prov. Park, 
Dow D-2A-C (v); Haney, Taylor 1002 (UBC); Hope-Prince Hwy., 
Brayshaw 49470 (uBC); Kelsey Bay, Calder & MacKay 29204 (WTU); 
Kitsault, Fisher (UBC); New Westminster, McCalla 2944 (ALTA); 
Prince Rupert, Mendel 78 (v); Prudhomme Lake, Taylor & Lewis 
618 (UBC); Terrace, Szczawinski (UBC, V). 

Kalmia microphylla var. occidentalis has been considered 
specifically distinct only by Small (1914). In all floristic 
works of the west coast of North America this taxon has 
been considered a part of the eastern K. polifolia, although 
it nas been rarely considered as a separate subspecies 
(Abrams, 1951). Hultén (1948) mentions that he was not 
able to separate the Alaskan specimens from those of East- 
ern America. He indicates, however, that a slight racial 
difference might exist, but that a large body of material 
would be necessary for a definite statement. 

Kalmia microphylla var. occidentalis and K. polifolia are 
strikingly similar. Both have the same general habit and 
size, and except for a slight difference in flower size these 
structures are also the same. Under close morphological 
examination, however, these two species are easily sepa- 
rated. In K. polifolia, the leaf margins are usually revolute, 
and well developed stalked glands are present on both sides 
of the midrib. In K. microphylla var. occidentalis, by con- 
trast, the leaves are not revolute, or only slightly so, and 
the stalked glands are absent. The two taxa also differ in 
seed size and calyx width. In K. polifolia, the seeds are long 

342 Rhodora [Vol. 76 

and narrow, 1.5-2.1 mm long, while the calyx varies in 
width from 5-7 mm. In K. microphylla var. occidentalis, 
the seeds are shorter and thicker, 1-1.4 mm long, and the 
calyx varies from 7-10 mm broad. 

The geographic distribution of the two taxa also differs, 
and they never come into contact with one another. Kalmia 
polifolia is mainly an eastern species, occurring as far west 
as Saskatchewan and northeastern Alberta, while K. micro- 
phylla var. occidentalis occurs along the west coast from 
southern Alaska, south through British Columbia and from 
Washington to Oregon. Most of the specimens of var. occi- 
dentalis examined were from marshes, bogs and wet open 
areas at lower elevations from coastal areas and the islands 
off the coast of Alaska and British Columbia. Hultén 
(1968), who did not distinguish between these two species, 
shows this difference in distribution and also has an excel- 
lent drawing of K. microphylla var. ? occidentalis (as K. 
polifolia subsp. polifolia) . 

Cytologically, Kalmia microphylla var. occidentalis (n = 
12) is more closely related to K. microphylla var. micro- 
phylla (n = 12) than it is to K. polifolia (n = 24). Crosses 
between var. occidentalis and K. polifolia result in triploid 
F, hybrids that produce less than 10% stainable pollen and 
no seeds (Jaynes, 1968a). In contrast, crosses between K. 
microphylla var. microphylla and var. occidentalis were 
highly fertile with the F,’s being intermediate between the 
two parents, producing viable seed and pollen that looked 
normal and fertile (Jaynes, persona] communication). 

Kalmia microphylla var. occidentalis f. alba Ebinger, 
forma nov. 

Forma typicae habita statura etc. similis, floribus omni- 
bus albis. 

This white flowering form, which is similar to that found 
in some other species of Kalmia, has been collected at two 
different localities. Alaska: near Wrangell, Mrs. J. C. Dart, 
1922. (us #1,118,961, Holotype). British Columbia: Lulu 
Island, peat bog between N. Westminster Highway and 
No. 4 road, H. Rhodes, et al., 29 May 1954 (UBC). 

1974] Genus Kalmia — Ebinger 343 

29. Kalmia polifolia Wang., Schr. Berlin Ges. Naturf. 
Freunde 8: 130. 1788. TYPE: unknown. 

Kalmia polifolia Ludwig, Neu. Wilde Baumz. 25. 1783. 
(nom. nud). 

Kalmia glauca Loddiges ex Ludwig, Neu. Wilde Baumz. 
25. 1783. (nom. nud.), Ait. Hort. Kew. 2: 64. 1789. TYPE: 
Plant growing in the Kew Gardens, (K). 

Chamaedaphne glauca | ( Ait.) Kuntze, Rev. Gen. Pl. 2: 
388. 1891. 

Kalmia glauca var. praecox Lavallée Arbor. Segrez. 159. 
1877. (nom. nud.). 

Kalmia glauca var. stricta Jager, Ziergeholze 279. 1865. 

Kalmia glauca var. superba Makoy ex Bosse, Vollst. 
Handb. Blumengart. 2: 348. 1841. (nom. nud.), Jager & 
Beissner, Ziergeholze 194. 1884. Kalmia superba K. Koch, 
Dendrologie 2(1): 154. 1872. (nom. nud.). 

Kalmia rosmarinifolia Dum.-Cours. Bot. Cult. 2: 250. 
1802. TYPE: unknown. Kalmia glauca p rosmarinifolia 
(Dum.-Cours.) Pursh, Fl. Am. Sept. 296. 1814. Kalmia 
polifolia var. rosmarinifolia (Dum.-Cours.) . Rehd. in Bailey, 
Cycl. Amer. Hort. 2: 854. 1900. 

Kalmia oleaefolia Dum.-Cours. Bot. Cult. ed. 2. 3: 322. 
1811. (as synonym). 

Sparingly branched shrub to 1 m tall; branchlets slightly 
flattened, 2-edged, glabrous or puberulent, commonly with 
decurrent ciliate lines from the petiole base; leaves oppo- 
site, subsessile with a broad petiole to 2 mm long, coriaceous, 
linear to oblong, the margins usually revolute, 1.5-4 cm 
long, 0.3-1.3 em broad, 214 to 4 times as long as broad, 
whitened beneath with small hairs, the midrib covered with 
purple clavate hairs on both surfaces; inflorescence a few- 
flowered, terminal corymbiform raceme, each flower on a 
glabrous pedicel 1.5-3 cm long, each pedicel subtended by 
a pair of bractlets in the axis of a leaf-like bract; calyx 
deeply 5-lobed, colorless to light pink, 5-7 mm broad, the 
lobes obtuse, margins ciliate ; corolla rose-purple, 1.2-1.8 cm 
broad; anthers purple, 1-1.5 mm long; fruit subglobose, 

344 Rhodora [Vol. 76 

4-7 mm broad, 4-6 mm long; seeds 1.5-2.2 mm long, the 
testa extending past the ends of the seed. Chromosome 
number n — 24, 

Distribution: Bogs, swamps, and other wet, open areas 
from northeastern Alberta east to the Atlantic Ocean and 

south to the Great Lakes, Pennsylvania, New York and 
New Jersey. 

Common name: Bog Laurel, Swamp Laurel, Pale Laurel, 
Gold Withy. 


UNITED STATES: Connecticut: HARTFORD CO.: Suffield, Harger 
6524 (ARIZ), Blewitt 3490 (PH). LITCHFIELD CO.: Bingham Pond, 
Weber 1489 (Isc). NEW HAVEN CO.: Middlebury, Shepard (conn). 
Maine: ANDROSCOGGIN CO.: Livermore, Gordon 657 (MAINE). AROOS- 
TOOK CO.: Knoles Corner, Neal 1908 (MAINE), True 6068 (PENN). 
CUMBERLAND CO.: Standish, Rossbach 5599 (ACAD, MAINE). FRANKLIN 
co.: Mt. Abraham, Rossbach 5389 (MAINE). HANCOCK CO.: Mt. 
Desert Isl, Redfield 4266 (US). KENNEBEC CO.: Monmouth, Hyland 
90 (MAINE). KNOX CO.: Rockland, Rossbach 1429 (ACAD). OXFORD 
co.: Old Spec, Richards 6032 (MAINE). PENOBSCOT CO.: Orono, 
Fernald 349 (GH, Mass, MIN, MSC, ND, NY, PH, US). PISCATAQUIS CO.: 
Greenville, True 4701 (PAC), 4814 (PENN). SOMERSET CO.: Jackman, 
True 4668 (PENN). WALDO CO.: Northport, Rossbach 2499 (ACAD), 
5349 (MAINE). WASHINGTON CO.: Jonesport, Steinmetz 844 (MAINE). 
YORK CO.: Sunken Pond, True 214 (GH, PENN). Massachusetts: 
BERKSHIRE C0.: Wolf Swamp, Walters (NHA). ESSEX CO.: Andover, 
Steele (MU). FRANKLIN CO.: Hawley, Forbes 24 (MASS). HAMPDEN 
co.: Ludlow, Seymour 557 (DUKE, GH, MASS, NY). HAMPSHIRE CO.: 
Belchertown, Torrey (MASS). MIDDLESEX CO.: Concord, Eaton 
(MAINE). WORCESTER C0.: Worcester, Dowling 818 (NY). Michigan: 
ALCONA CO.: Hall & Hagenah 447 (BLH). ALPENA co.: Hatt (BLH). 
BARAGA CO.: Baraga, Richards 1783 (MICH, NY), 2956 (MAINE, MICH). 
CHARLEVOIX CO.: Beaver Isl, Reis 906 (BLH, MSC). CHEBOYGAN CO.: 
Mud Lake, Gleason, Jr. 665 (DUKE, GH). CHIPPEWA CO.: Sugar Isl., 
McVaugh 8745 (BLH, CAN, MICH). CLARE CO.: Lake Station, Dreis- 
bach 5650 (MICH). CRAWFORD Co.: Hartwick, Bingham (BLH). 
DELTA CO.: Bull Run Creek, Hagenah 6723 (BLH). EMMET CO.: Carp 
Lake, Gates & Gates 10334 (RM). GOGEBIC CO.: Watersmeet, Voss 
7846 (MICH), 12472 (MICH). GRAND TRAVERSE CO.: "Traverse City, 
Dieterle 1299 (cM, MICH). GRATIOT C0.: Alma, Davis (MONT). 
HOUGHTON CO.: Laurium, Herman 7591 (NY, US). IRON CO.: Golden 
Lake, Voss 4122 (MICH), KALKASKA CO.: Carlisle, Hagenah 4602 
(BLH). KEWEENAW CO.: Mott Isl, Bailey & Bailey 4131 (MICH). 

1974] Genus Kalmia — Ebinger 345 

LEELANAU CO.: Maple City, Thompson L-1693 (BLH). LENAWEE CO.: 
Clinton, Wood & Wood (us). LUCE CO.: Spring Lake, Hagenah 1913 
(BLH). MACKINAC CO.: Hagenah 4876 (BLH). MANISTEE CO.: Manis- 
tee, Daniels 4267 (MONT). MARQUETTE CO.: Turin, Barlow (NY). 
MIDLAND CO.: Sanford Game Preserve, Dreisbach 7941 (MICH). 
MONTMORENCY CO.: Barding Swamp, Case (MICH). OAKLAND CO.: 
Lakeville, Farwell 4905 (BLH). OGEMAW CO.: Clear Lake, Case, Jr. 
(MICH). ONTONAGON CO.: Berglund, Beaman 1848 (MS). OSCEOLA 
co.: Evart, Monaghan (MSC). OSCODA CO.: Zimmerman 416 (MICH). 
OTSEGO COo.: Otsego Lake Village, Voss 10847 (MICH). ROSCOMMON 
co.: Higgins Lake, Denton 1038 (MSC). ST. CLAIR CO.: Capae, Dodge 
(MU, TENN). SANILAC CO.: Minden City, Voss 8341 (MICH). SCHOOL- 
CRAFT CO.: Long Lake, Voss 2657 (MICH). WEXFORD CO.: Cadillae, 
House 4660 (Us). Minnesota: BELTRAMI CO.: Star Isl, Butters & 
Rosendahl 6584 (MIN). CARLTON CO.: Atkinson, Lakela 3550 (DUL, 
MIN). CASS CO.: Lake Kilpatrick, Ballard (ORE, RM, US, WS). CLEAR- 
WATER CO.: Long Lake, Morley 1106 (MIN). COOK CO.: Sea Gull 
Lake, Lakela 3669 (DUL, MIN). CROW WING CO.: Bay Lake, Sheldon 
2172 (MIN). ITASCA CO.: Sand Lake, Johnson 2042 (PH), 2207 (GH, 
US). KANABEC C0.: Mora, Moore & Tryon, Jr. 17504 (MIN). KOOCHI- 
CHING Co.: Rainy Lake, Moore & Moore 11793 (ISC, MIN). LAKE 
co.: Basswood Lake, Lakela 8898 (DUL). PINE CO.: Bruno, Jerabek 
66 (MIN). ST. LOUIS CO.: Palo, Nickelson 2432 (ISC, NY, ws). New 
Hampshire: BELKNAP CO.: Crystal Lake, Hodgdon & Steele 9125 
(NHA). CARROLL CO.: Church Pond, Gale (NHA). CHESHIRE CO.: 
Keene, Gilbert (NY, PH). coos Co.: Mt. Washington, Hartmann 
207 (NY); Moore 260 (Gv), 1081 (GY). GRAFTON CO.: Mt. Lafayette, 
Fernald & Smiley 11839 (GH). HILLSBORO CO.: Peterborough, Batch- 
elder 1891 (KY). ROCKINGHAM CO.: Derry, Batchelder (NHA). 
STRAFFORD CO.: Barrington, Hodgdon & White 11935 (NHA). SULLI- 
VAN Co.: Springfield, Hodgdon & Steele 9128 (NHA). New Jersey: 
MORRIS CO.: Mt. Hope, Edwards & Rusling (CHRB). SUSSEX CO.: 
Kittatinney Ridge, Griscom 9562 (GH). New York: CHAUTAUQUA CO.: 
Petit, Thurber (GH). COLUMBIA CO.: Knickerbocker Lake, McVaugh 
1013 (PENN). ERIE CO.: Protection, Miller 3764 (MSC). ESSEX CO.: 
Whiteface Mt., Rudd 824 (US). FRANKLIN C0.: Upper Savanac Lake, 
Guest 182 (PAC, PENN). HAMILTON CO.: Bern Bog, Henry (CM). 
HERKIMER CO.: Twin Lake, VanPelt & Long (PH). MADISON CO.: 
Recksport, House 6930 (US). ONEIDA CO.: White Lake, True 38 
(PENN). ONONDAGA CO.: Labrador Pond, Clausen 2129 ANY, PAC, 
PENN). OSWEGO CO.: Mud Pond, Wiegand 13764 (GH). ST. LAWRENCE 
co.: Norfolk, Phelps 757 (GH, NY, US). SARATOGA CO.: Saratoga 
Lake, House 27663 (PENN). SCHENECTADY CO.: Princetown, Svenson 
(GH). SULLIVAN CO.: Neversink, Jones, Phillips & Stebbins 21 (GH). 
TIOGA CO.: Owego, Mapes 4767 (GH). ULSTER CO.: Rockland, Camp 
3036 (NY). WASHINGTON CO.: Salem, Stewart 63 (NY). Pennsylvania: 

346 Rhodora [Vol. 76 

CARBON CO.: Albrightsville, Pretz 13321 (PH). LACKAWANNA CO.: 
Elmhurst, Glowenke 4820 (PENN). MONROE CO.: Tannerville, Leeds 
2237 (MIN, PH). PIKE CO.: Shohola Falls, Byhouwer & Kobuski 226 
(GH). SUSQUEHANNA Co.: Ararat, Glowenke 184 (PENN). WAYNE 
co.: Lehigh Pond, Adams 3699 (GH). Rhode Island: PROVIDENCE CO.: 
Glocester, Collins (CONN, GH, MASS, NHA, NY, PENN, PH). Vermont: 
ADDISON CO.: Ripton, Eggleston 280 (GH). CALEDONIA CO.: Peacham, 
Blanchard (PH, US). CHITTENDEN CO.: Colchester, Pringle (BRY). 
LAMOILLE CO.: Mt. Mansfield, Moseley (MASS). ORLEANS CO.: Lake 
Willoughby, Stevens (US). RUTLAND CO.: Mt. Tabor, Seymour 24407 
(BRY). WINDHAM CO.: Tyson, Moldenke & Moldenke 10263 (Ny). 
Wisconsin: BAYFIELD C0.: Sand Bay, Cheney 6371 (UWM). DOOR CO.: 
Bailey’s Harbor Ridge, Shinners (UWM). FLORENCE CO.: Spruce 
Lake, Mau (UWM). FOREST CO.: Crandon, Palmer 27776 (GY). IRON 
co.: Mercer, Salamun 228 (UWM). JACKSON CO.: Ketchum, Hartley 
& Morrissey 9158 (ISC). KEWAUNEE CO.: Gregor (OS). LINCOLN CO.: 
Tomahawk, Engel (UWM). MARINETTE CO.: Walsh, Throne (UWM). 
MONROE CO.: La Grange Twp., Hartley 7884 (MIN). ONEIDA CO.: 
Three Lakes, Wadmond 2758 (MIN). RUSK CO0.: Ladysmith, Goessl 
2162 (UWM). SHAWANO CO.: Camp Tivol, Keefe (UWM). TAYLOR 
co.: Buch Lake Area, Piehl (UWM). viLAS co.: Eagle River, 
Hollister 334 (US). WASHBURN CO.: Spooner, Cheney 6135 (UWM). 

CANADA: Alberta: Lake Athabaska: Smith Landing, Preble & 
Preble 99a (US); Alta, Raupp & Abbe 4529 (ALTA, CAN, MIN, PH). 
Gregoire Lake, Dumais & Anderson 2693 (ALTA); Richardson Lake, 
Whitehorn & Barber 185 (SASK). Labrador: Anatolak, Sewall 469 
(GH, US), 471 (GH, US); Ashuanipi Lake, Harper 3877 (CAN, MIN, 
us); Attikamagen Lake, Harper 3501 (CAN, ISC, MIN, PENN); Battle 
Harbour, Koelz 36 (MICH, US); Cape St. Charles, Potter & Brierly 
3410 (GH, NMC); Cartwright, Hitchcock 23818 (vs); Gannet Isl, 
Potter & Brierly 3411 (GH); Goose Bay, Gillett & Findley 4931 
(ACAD, NY); Gready Isl., Bishop 474 (GH), 475 (GH); Hopedale, 
Walker 370 (PENN); Mealy Mts., Gillett & Findley 5273 (ACAD, 
NY, TRT, US); Mills Harbour, Cook 88 (GH, US); Petty Harbour, 
Bishop 476 (CAN, GH); Spotted Isl, Wallace 42 (CONN); Sandwich 
Bay, Woodworth 329 (GH), 330 (GH); Windy Tickle, Potter & Brierly 
3407 (MASS). Manitoba: Baralzon Lake, Scoggan & Baldwin 8216 
(GH, MIN, WIN); Cranberry Portage, Freedmann (WIN); Churchill 
Dist., Schofield 1001 (DUL, GH, MIN, WS); Duck Lake, Scoggan & 
Baldwin 8245 (ALTA, CAN, GH); Gillam, Punter 97 (WIN); Goldesand 
Lake, Keleher 107 (WIN); Heming Lake, Keleher 159 (WIN); Herch- 
nier, Core 215 (WVA); Lake Winnipeg, Lowe (WIN); Lake Winni- 
pegosis, Scoggan & Baldwin 8152 (CAN); Nelson River, Scoggan 
3016 (ALTA, CAN, MIN, WIN); Nueltin Lake, Baldwin 2180 (CAN, TRT, 
WIN, WS); Oxford Lake, Scoggan 5390 (CAN); Reindeer Lake, Bald- 

1974] Genus Kalmia — Ebinger 347 

win 2028 (CAN, TRT, WIN); Seal River, Ritchie 1794 (CAN, MIN); 
Third Cranberry Lake, Freedman (WIN); Wabowden, Gillett 2790 
(us); Whiteshell Forest Reserve, Scoggan 8781 (ALTA, WIN); York 
Factory, Punter 134 (WIN). New Brunswick: ALBERT CO.: Grindstone 
Isl, Christie 1576 (NBM). CHARLOTTE CO.: Campobello Isl., Malte 
955/29 (CAN). GLOUCESTER CO.: Bathurst, Malte 13 (CAN). KENT 
co.: Little Forks Brook, Squires & Squires (NBM). MADAWASKA CO.: 
St. Leonard, Maite 605 (CAN). QUEENS CO.: Castaway, Gorham & 
Squires (NBM). SAINT JOHN CO.: Portland, Matthew (NBM). WEST- 
MORELAND CO.: Sackville, Fowler (QUK). YORK CO.: Fredericton, 
Dore & Gorham 4569 (ACAD). Newfoundland: BONAVISTA SOUTH DIST.: 
Lethbridge, Rouleau 8363 (NFLD). BURGEO-LA POILE DIST.: Grandy 
Brook, Fernald, Long & Fogg 359 (GH, PENN, PH). FERRYLAND DIST.: 
Witless Bay, Rouleau 5391 (CAN, NFLD, US). FOGO DIST.: Fogo Isl., 
Sornborger (GH). FORTUNE BAY-HERMITAGE DIST.: Hermitage Bay, 
Palmer 1353 (GH, US). GRAND FALLS DIST.: Bishop Falls, Fernald & 
Wiegand 6023 (CAN, GH, K, NY, PH). HUMBER DIST.: Eagle's Nest 
Brook, Rouleau 3069 (NFLD). ST. BARBE DIST.: Trout R., Rouleau 
3515 (NFLD). ST. MARY'S DIST.: Colinet, Rouleau 4175 (NFLD). ST. 
JOHN’S DIST.: Doctor Hill, Fernald, Long & Fogg 1916 (GH, PENN, 
PH, US). TRINITY NORTH DIST.: George’s Brook, Rouleau 6078 (NFLD). 
WHITE BAY DIST.: Twin Lakes, Rouleau 6525 (NFLD, US). Northwest 
Territories: Baralzo Lake, Scoggan & Baldwin 8335 (CAN), 8379 
(CAN); Gagnon Lake, Scotter 3162 (CAN); Ingalls Lake, S.C.Z. 
(CAFB). Nova Scotia: ANNAPOLIS CO.: Karsdale, Johnson (ACAD). 
CAPE BRETON CO.: Scatari Isl, Ershine 51-1167 (QUK). COLCHESTER 
co.: Folleigh Lake, Perry (ACAD). CUMBERLAND CO.: Sand R., 
Schofield & Webster 5592 (ACAD). DIGBY CO.: Central Grove, Fernald 
& Long 24285 (GH). GUYSBOROUGH CO.: Tor Bay, Eshbaugh 522 
(IND). HALIFAX CO.: Fernald, et al. 24284 (GH, PH). HANTS CO.: 
Mt. Uniacke, Jack 3230 (GH), 3231 (GH). INVERNESS CO.: Mackenzie 
Mt., Rossbach 4235 (ACAD). KINGS CO.: Caribou Bog, Erskine 
(ACAD). LUNENBURG CO.: Deep Cove, Smith, et al. 19604 (ACAD, 
QUK). PICTOU co.: New Glasgow, Dore & Gorham 45-389 (ACAD). 
QUEENS C0.: Eagle Head, Donly 1009 (ACAD). RICHMOND CO.: Grand 
R., Smith, et. al. 10110 (ACAD, TRT). SHELBURNE CO.: Barrington, 
Fernald, Long & Linder 22154 (CAN, GH). VICTORIA CO.: Cape Breton 
Nat’l. Park, Armstrong 452 (QUK). YARMOUTH CO.: Yarmouth, 
Pease & Long 22152 (GH, PENN). Ontario: ALGOMA DIST.: Michi- 
picoten Harbour, Hosie, Harrison & Hughes 1223 (CAN, TRT). 
CARLETON CO.: Mer Blue, Porsild 7743 (CAN). COCHRANE DIST.: 
Fraserdale, Kirk 75 (TRT): Timmins, Baldwin 4761 (CAN, TRT). 
DURHAM C0.: Newtonville, Krug & Purchase 473 (TRT). FRONTENAC 
co.: Kingston, Beschel 10128 (QUK). GREY CO.: Euphrasia Twp., 
Jackson (TRT). HALIBURTON CO.: Dorset, Walshe 12 (CAN, TRT). 
HURON C0.: Wingham, Anderson 724 (TRT). KENORA DIST.: Rich- 

348 Rhodora [Vol. 76 

mond Gulf, Abbe & Abbe 3383 (CAN, MIN); Las Seul, Baldwin 8739 
(CAN, MIN, QUK, TRT); Big Trout Lake, Moir 13 (MINN), 21 (CAN, 
MIN), 1744 (MIN); Swan Lake, Moir 156 (MIN); Fort Severn, 
Moir 1337 (CAN, MIN); Black Duck R., Moir 1831 (MIN), 2037 (CAN, 
MIN, TRT) ; Windigo R., Cringan P-26 (ACAD, TRT). MANITOULIN DIST.: 
Great Duck Isl, Grassel 6028 (NY). MIDDLESEX CO.: London, Baker 
1638 (OAC). MUSKOKA DIST.: Huntsville, Soper & Shields 4927 
(TRT), 4967 (TRT); Margaret Lake, Chow 6415 (TRT). NIPISSING 
DIST.: Algonquin Park, Watson 3967 (TRT). PARRY SOUND DIST.: 
Axe Lake, Shields & Shields 1640 (TRT). PEEL CO.: Heart Lake, 
Catling & Sparling (TRT). RUSSELL CO.: Bourget, Heimburger 2612 
(TRT). SUDBURY DIST.: Skead, Winterhalder 66114 (CAN). THUNDER 
BAY DIST.: Bear Trap Lake, Voss 10361 (MICH, TRT). TIMISKAMING 
DIST.: Hilliard Twp., Baldwin & Breitung 2564 (CAN, SASK, TRT). 
WATERLOO CO.: Baden, Montgomery 997 (au, OAC). WELLAND CO.: 
Humberstone, Soper & Dale 3776 (GH, MIN, TRT, US). WELLINGTON 
Co.: Puslinch, Stroud (TRT). YORK co.: Vandorf, Robb (TRT). 
Prince Edward Island: KING Co.: Bothwell, Fernald & St. John 11148 
(CAN, GH, US, WS). PRINCE CO.: North Point, Erskine & Dore 1080 
(ACAD, NY). QUEENS CO.: Brackley Point Rd., Fernald, Long & St. 
John 7901 (CAN, GH, PH, US, WS). Quebec: ABITIBI CO.: Watson 
Lake Area, Bentley 58145 (TRT), 5837 (CAN, TRT). BERTHIER CO.: 
Lanoraie, Scoggan 1538 (CAN). BONAVENTURE CO.: Nouvelle, Collins 
& Fernald (GY). CHAMBLY CO.: Saint-Bruno, M.-Victoria & R.- 
Germain 43884 (cM, GH, ND, PH). CHICOUTIMI CO.: Lac Merlac, 
Brisson 63124 (CAN). GASPE WEST CO.: Mt. Albert, Scoggan 1086 
(CAN). GATINEAU CO.: Eardley, Cody & van Rens 11167 (DUKE, GH, 
MSC, NY, TENN). HULL CO.: King Mt., Zavitz 1107 (TRT). HUNT- 
INGTON CO.: Covey Hill, M.-Victoria & R.-Germain 43300 (GH). 
LABELLE C0.: Nominingue, Roy 1167 (wvA), 3650 (ACAD, CONN, NY, 
RM, US). LAKE ST. JOHN WEST CO.: Peribonka, M.-Victorin 15849 
(GH). LEVIS CO.: St. Nicolas, Cayouette & Cayouette 54-89 (TRT). 
MAGDALEN ISLANDS CO.: Grindstone Isl, Fernald, et. al. 7900 (CAN, 
PH, WS). MATANE CO.: Montagne Lake, Boivin 787 (PENN). MATA- 
PEDIA CO.: Sayabee, Scoggan 13600 (CAN). MEGANTIC CO.: Black 
Lake, Fernald & Jackson 12137 (GH). MONTCALM CO.: Lac Tapani, 
Pennell 16686 (PH). MONTMAGNY CO.: Lake Monroe, R.-Germain & 
Courtemanche 454 (US). PAPINEAU CO.: Notre Dame du Laus, 
C.-Joseph 8508 (ACAD). PONTIAC COo.: Danford Lake, Dore 19994 
(TRT). QUEBEC CO0.: Montmorency Fall, Macoun 68560 (am). 
RIMOUSKI CO.: Father Point, Bartram & Long 581 (PH). SAGUENAY 
co.: Fort Chimo, Porsild 21873 (CAN); Knob Lake, Harper 3350 
(HNH, QUK); Cairn Isl., Abbe, et. al. 3067 (RM): Seven Isl., Robinson 
868 (CAN, GH, NY); Mutton Bay, Lewis 582 (CAN). ST. JEAN CO.: 
Villeneuve, M.-Victorin, et. al. 43047 (MU). SHERBROOKE CO.: Orford, 
Forest 16058 (CAN, CM, QUK, SASK). TERREBONNE CO.: St. Janvier, 

1974] Genus Kalmia — Ebinger 349 

Scoggan 314 (CAN). TWO MOUNTAINS CO.: Oka, M.-Victorin 8228 
(NY, US). WOLFE C0.: Lac Aylmer, Hamel 13217 (CAN). Saskatche- 
wan: Athabaska Lake, Lang 188 (Us); Raupp 6175 (CAN, GH, NY), 
6657 (GH), 6731 (GH, NY); Bittern Cr., Wheaton (CAFB); Candle 
Lake, Boivin & Breitung 6157 (NY); Cree Lake, Maini 21 (RM, 
SASK), 235 (SASK), 236 (CAN, RM, SASK); Crooked R., Breitung 70 
(SASK); Cumberland House, Argus 4083 (SASK); Hatchet Lake, 
Maini 91 (OAC, RM, SASK); Nipawin Prov. Park, Argus 4122 (SASK); 
Orley, Breitung 268 (ALTA, CAN, NY, US); Prince Albert Nat’l. Park, 
Fraser (PAC, SASK); Stoney Rapids, Maini 205 (RM, SASK), 275 
(SASK), 462 (RM, SASK); White Fox, Fraser (SASK) ; Windrum Lake, 
Bryenton 166 (CAN). 

The name Kalmia polifolia was first used by Ludwig 
(1783), who gave it the common name Canada Laurel or 
Poley Leaved Laurel, but did not describe it. This name 
first appears with a valid description a few years later 
(Wangenheim, 1788). He gives its distribution as extend- 
ing northward from 45 degrees latitude in America, Nova 
Scotia and the colder parts of Canada. His description, 
illustration, and the distribution leave no doubt as to the 
taxon referred to. Recently, Boivin (1967) suggested that 
the name be spelled poliifolia to conform to article 73 of the 
International Code of Botanical Nomenclature (Lanjouw, 
1966). The name K. glauca was first used as a synonym 
of K. polifolia by Ludwig (1783) and later described by 
Aiton (1789). The type was based on a plant from New- 
foundland growing at Kew. A specimen in the Kew Her- 
barium has written on the label “Kalmia glauca Hort. 
Kews. sp. nov. 1784” and probably represents the type of 
this name. Kalmia rosmarinifolia was used by Dumont de 
Courset (1802) to distinguish individuals that have narrow 
leaves with strongly revolute margins. Later, he listed the 
name as a synonym of K. glauca (Dumont de Courset, 

Kalmia polifolia is found in bogs, swamps, and other wet 
places, commonly forming a border around ponds and 
lakes. In bogs, its roots usually form dense mats that 
extend out over the water. It is also found at higher ele- 
vations in the mountains of northeastern United States and 
Canada. This species is the most widely distributed member 

350 Rhodora [Vol. 76 

of the genus, extending from northeastern Alberta, across 
Canada to the east coast and south to New Jersey. Its 
entire range, however, is hard to determine with certainty. 
There are two reports of its occurrence as far north as the 
Arctic Circle in the Northwest Territories (Gussow, 1933, 
and Thieret, 1963). Both of these (Maguse Lake, Gussow 
33; Contwoyto Lake, Thieret 9415) are specimens of K. 
microphylla var. microphylla. The only specimens of K. 
polifolia seen from the Northwest Territories are from the 
extreme southern part (Scotter, 1966). In the Great Lakes 
region this taxon extends as far south as northern Illinois 
(Fuller, 1944), and along the east coast to New Jersey 
(Griscom, 1931). This species has also become naturalized 
in Europe since Scott (1962) reported that it has been col- 
lected from Flanders Moss, Perthshire, Scotland. 

Besides leaf size and shape, the character most commonly 
used by previous authors to separate Kalmia polifolia from 
K. microphylla is revolute leaf margins. In K. microphylla, 
the leaf margins are not revolute, or only slightly so, while 
in K. polifolia the margins are usually strongly revolute. 
This characteristic is somewhat variable. In about 80% of 
the specimens examined, the leaf margins were strongly 
revolute, resulting in a narrow, linear leaf. In the remain- 
ing specimens, some lacked revolute margins entirely, while 
in others the younger leaves were revolute while older 
leaves were not. The variability of this characteristic is 
responsible, in part, for the confused taxonomic status of 
K. microphylla. 

The major distinguishing vegetative character that sep- 
arates Kalmia polifolia from the related species, K. micro- 
phylla, is the presence of purple clavate hairs on the leaf 
midrib. Boivin (1968) appears to be the first to have made 
use of this character when he separated variety microphylla 
from variety polifolia. He mentions, however, that west of 
Alberta the distinction of the two varieties becomes mean- 
ingless, as most of the British Columbia material is inter- 
mediate, the leaves being long and narrow like variety poli- 
folia, but glandless on the midrib. It appears that he was 

1974] Genus Kalmia — Ebinger 851 

confusing K. polifolia with the western K. microphylla var. 
occidentalis, which is similar vegetatively, except for the 
glandless midribs. Clavate glands were found on all speci- 
mens of K. polifolia examined. These glands vary in size, 
shape and color, and are more plentiful on some specimens 
than on others. The glands average about 0.3 mm long, are 
colorless near the base and become deep purple to black 
toward the swollen apex. In some specimens, however, they 
are nearly colorless throughout, and in others barely ex- 
panded above. Also, their distribution varies, being more 
common on some leaves than on others. On older leaves, it 
is not uncommon to find that many of the glands have 
broken off. The glands on the upper side of the midrib are 
commonly shorter, darker colored and not as obvious as 
those on the under side of the leaf. For determining the 
presence of these glands it is best to look at the developing 

Morphologically and genetically Kalmia polifolia (n = 
24) is most closely related to K. microphylla (n = 12). The 
F, of a cross between these two species is a highly viable 
triploid which has a pollen stainability of 0-15% (Jaynes, 
1968a). These similarities suggest that these two taxa have 
probably become specifically distinct as a result of the last 
ice age and the resulting development of a tetraploid taxon 
from a wide ranging ancestral diploid species. Jaynes 
(1969) suggested that these taxa should be considered sep- 
arate species based on his cytological and genetic studies. 
Kalmia polifolia can also be crossed with other members of 
the genus, but few viable offspring are produced. 

Kalmia polifolia forma leucantha Schofield & Smith, 
Canad. Field Nat. 67: 94. 1953. TYPE: Newfoundland: 
near Hodgewater Line, Trinity South, E. E. Smith and 
A. C. Smith 1055 (ACAD). i 

This white flowering form, which is similar to that found 
in some of the other species of Kalmia, is known from one 
collection. This single specimen was found in a bog where 
the typical form was very common. 

352 Rhodora [Vol. 76 

3. Kalmia latifolia Linnaeus, Sp. Pl. 1: 391. 1753. 
TYPE: “Habitat in Marilandia, Virginia, Pennsylvania" 
(LINN 560.1, lectotype, specimen at the top right). 

Chamaedaphne latifolia (L.) Kuntze, Rev. Gen. Pl. 2: 
388. 1891. 

Kalmia latifolia var. acuminata Raf. Medical Fl. 2: 17. 
1830. (nom. nud.). 

Kalmia latifolia var. alba Raf. Medical Fl. 2: 17. 1830. 
Kalmia latifolia f. alba (Mouillefert) Rehd. Rhodora 12: 2. 

Kalmia latifolia var. arborea Raf. Medical Fl. 2: 17. 1830 
(nom. nud.). 

Kalmia latifolia var. pavarti André, Revue Hort. 60: 541. 

Kalmia latifolia f. rubra K. Koch, Dendrologie 2(1) : 152. 
1872. Kalmia latifolia var. rubra (K. Koch) Rehd. in 
Bailey, Cycl. Amer. Hort. 2: 854. 1900. 

Kalmia latifolia var. ternata Raf. Medical Fl. 2: 17. 1830. 

Kalmia latifolia var. laevipes Fern. Rhodora 42: 53. 1940. 
TYPE: Virginia: NORFOLK CO.: dry woods of a *hammock", 
Great Dismal Swamp, west of Yadkin, Fernald & Long 
11101 (GH). 

Kalmia lucida K. Koch, Dendrologie 2(1): 152. 1872. 
(as synonym). 

Kalmia nitida Forbes, Hort, Woburn. 93. 1833. TYPE: 
unknown, probably a living plant. 

Much branched shrub to 4 m tall (rarely a small tree) ; 
branchlets terete, reddish brown, becoming gray, stipitate- 
glandular to glabrous; leaves alternate (appearing nearly 
whorled on slow growing twigs), flat, coriaceous, elliptic to 
elliptic-lanceolate, 4-12 cm long, 1.5-4 (rarely 5) em wide, 
dark green and glabrous above, light green to reddish brown 
and with short glandular hairs beneath, apex acute, base 
narrowly cuneate, midrib puberulent above; petiole 1-3 em 
long, glabrous to puberulent, sometimes glandular-stipitate; 
inflorescence a terminal, compound corymb; pedicels 2-4 cm 
long, puberulent to stipitate-glandular, often floccose, viscid 

1974] Genus Kalmia — Ebinger S00 

(rarely glabrous), subtended by 2 bracteoles in the axil of 
a puberulent, lanceolate bract, 1-5 mm long; major inflor- 
escence branches puberulent, usually stipitate-glandular, to 
4 cm long; calyx glabrous to stipitate glandular, green to 
reddish, 5-8 mm broad, the lobes usually oblong, acute; 
corolla usually light pink (white to deep red) with purple 
spots around each anther pocket, usually lightly stipitate- 
glandular outside, puberulent inside, 1.5-3 cm broad; style 
1-1.8 cm long; anthers dark purple to brown, 1.2-1.7 mm 
long; filaments puberulent; capsule depressed globose, stipi- 
tate-glandular, 4-7 mm broad, 3-5 mm long; seeds .7-1 mm 
long, the testa loose and extending slightly past the ends of 
the seed. 

Distribution: Mostly rocky or sandy hardwood forests in 
eastern United States from southern Maine, west through 
southern New York to central Ohio, south to southern Mis- 
sissippi, Alabama, and Georgia and northwestern Florida. 

Common Name: Mountain Laurel, Broad-leaved Laurel, 
Calico-bush, Spoonwood, Ivy, Mt. Ivy, Big-leaved Ivy, 
Laurel-leaves, Calmoun. 


Alabama: CALHOUN CO.: Pendergrass (TENN). CHEROKEE CO.: 
Centre, Clonts 95 (1sC). CHOCTAW CO.: Silas, McDaniel 4218 (VDB). 
CLARKE CO.: Tallahatta Creek, Kral 19749 (VDB). CLEBURNE CO.: 
Kral 31843 (VDB). CONECUH CO.: Castleberry, Howell 760 (us). 
CULLMAN CO.: Cullman, Kral 28911 (VDB). DALE CO.: Newton, God- 
frey 55477 (TENN). DE KALB CO.: Mentone, Demaree 50521 (TENN). 
ESCAMBIA CO.: Little Escambia River, Ahles 7229 (MISSA). FRANK- 
LIN CO.: Russellville, Kral 26321 (VDB). HENRY co.: Headland, 
Wiegand & Manning 2370 (CU). JACKSON co.: Flat Rock, Kral 
34948 (VDB). LEE CO.: Chewacla, Barnes 119 (VDB). MARION CO.: 
Hackelberg, Kral 20024 (VDB). MARSHALL CO.: Albertville, Harbison 
15887 (TENN). MOBILE CO.: Mt. Vernon, Burnham 84 (VDB). PIKE 
co.: Troy, McDaniel 7830 (vpB). TUSCALOOSA Co.: Lake Nichols, 
Williams 104 (VDB). WASHINGTON CO.: Bates Creek, Segars 191 
(WVA). WINSTON cCOo.: Haleyville, Kral 29476 (vpB). Connecticut: 
FAIRFIELD CO.: Wilton, Eames 10426 (CONN). HARTFORD CO.: Wind- 
sor, Ahles 65351 (MASS). LITCHFIELD CO.: Preston, Lumsden (MU). 
MIDDLESEX CO.: Millers Pond, North 51 (MICH, US). NEW HAVEN CO.: 
Milford, Fames 2813 (CONN). NEW LONDON CO.: Lantern Hill, Glea- 

354 Rhodora [Vol. 76 

son (DUKE). TOLLAND CO.: Andover, Travis 2408 (CONN). WINDHAM 
co.: Crystal Lake, Peterson (cU). Delaware: NEW CASTLE CO.: 
Arden, Travis 1086 (PENN). SUSSEX CO.: Gumboro, Beaven 241 
(DUKE). Florida: ESCAMBIA CO.: McDavid, Goodale (MASS). GADSDEN 
co.: Sycamore, Godfrey 61925 (Msc). HOLMES CO.: Darlington, 
Godfrey 55498 (TENN). LIBERTY CO.: Greensboro, Karl 1916 (TENN). 
OKALOOSA CO.: Laurel Hill, Godfrey 61284 (US). WALTON CO.: De 
Funiak Springs, Perkins & Hall 2510 (cv). Georgia: CLARK CO.: 
Middle Oconee River, Duncan & Duncan 4358 (ID, IND, ISC, PENN, 
RENO, TENN, US). CLAY CO.: Cemochechahee Creek, Thorne 3453 
(CU). DADE CO.: Lookout Mt., MeVaugh 9023 (MICH), EARLY CO.: 
Sawhatchee Creek, Duncan & Thorne 1778 (CU, TENN). EFFINGHAM 
co.: Savannah River, Harper 1814 (US). GWINNETT CO.: McGuires 
Mill, Small (US). MORGAN CO.: Hard Labor Creek St. Park, Jones 
15456 (VDB). PICKENS CO.: Jasper, Cronquist 4483 (NO, US). RABUN 
cc.: Darling Spring, J.H.M. E1525 (WVA). RANDOLPH CO.: Cuthbert, 
‘Harbison 13729 (TENN). SCREVEN CO.: Newington, Cronquist 5023 
(MICH, US, WS). SUMTER CO.: Americus, Harper 523 (US). UNION 
co.: Vogel St. Park, McVaugh & Pyron 841 (ORE). UPSON co.: Flint 
River, Browne & Browne, Jr. 6587 (KY). WALKER CO.: Pigeon Mt., 
Wilson 176 (US). WALTON Co.: Athens, Cronquist 4468 (MICH, NO, 
US). WHITE CO.: Cleveland, Lovett (DUKE). WHITFIELD CO.: Dalton, 
Cronquist 5045 (MICH, US, WS). Indiana: CLARK CO.: Borden, Deam 
24495 (IND, MIN). CRAWFORD CO.: Riddle, Deam 16497 (IND). PERRY 
co.: Derby, Ramsey 10876 (IND). Kentucky: BELL co.: Pike Mt. 
St. Park, Collins 180 (KY). BREATHITT CO.: Robinson Forest, Smith 
1726 (KY). CARTER CO.: Cascade Cavern, Gilbert & McCoy 93513 
(us). CASEY CO.: Liberty, Wharton 4366 (KY). EDMONSON CO.: 
Mammoth Cave Nat’l. Park, Pittillo 2452 (Kv). ESTILL CO.: South 
Irvine, Wharton 2298b (Kv, MICH). FLEMING CO.: Plummer's Mill, 
Wharton 3873 (KY). FLOYD CO.: Hueysville, Centers & Blair TT 
(KY). GRAYSON CO.: Clifty Creek, Browne & Browne, Jr. 6494 (KY). 
HARLAN CO.: Poor Fork, Kearney 182 (ISC, MSC, OS, US). JESSAMINE 
co.: Camp Daniel Boone, Webb 104 (Ky). KNOTT CO.: Littcarr, 
Smith 59 (KY). LETCHER CO.: Browne & Browne, Jr. 5400 (KY). 
LEWIS CO.: Vanceburg, Wharton 5012 (KY, MICH). LINCOLN CO.: 
Ottenheim, Wharton 4866 (KY, MICH). MCCREARY CO.: Cumberland 
Falls, Rogers 55 (DUKE, IND, MIN, PENN, TENN, US, WVA). MADISON 
co.: Berea, Nelson (MICH, MIN, MU). MENIFEE CO.: Frenchburg, 
MelInteer 2302 (KY). MORGAN CO.: Ezel, Oldfield 57 (KY). POWELL 
co.: Stamping Ground, Singer (BRY). PULASKI CO.: Mt. Victory, 
Warden 129 (Kv). TODD CO.: Clifty, McCoy (WVA). TRIMBLE CO.: 
Bedford, Born (KY). WOLFE CO.: Sky Bridge, McInteer 1055 (KY). 
Louisiana: WASHINGTON PARISH: Bogalusa, Cocks (NO); Varnodo, 
Thieret 23481 (DUKE). 

Maine: CUMBERLAND CO.: Richville, Hyland 444 (MAINE). OXFORD 

1974] Genus Kalmia — Ebinger 355 

co.: Fryeburg, Hyland 446 (MAINE). PENOBSCOT CO.: Carmel, 
Harvey 622 (US). SAGADAHOC CO.: Phippsburg, Gordon 592 (MAINE). 
WASHINGTON CO.: Cherryfield, Hyland 760a (MAINE). YORK CO.: 
Kennebunk, Hyland 252 (MAINE). Maryland: ALLEGANY CO.: Cum- 
berland, Richards 128 (MAINE). ANNE ARUNDEL CO.: Camp Whip- 
poorwil, Grimes (WVA). BALTIMORE C0.: Abingdon, Nieuwland 
(ND). CAROLINE CO.: Newton, Quidas 37 (MAINE, PENN). CECIL CO.: 
Elkton, Randolph & Randolph 124 (CU). FREDERICK Co.: Hunting 
Creek, Earle 1872 (PENN). GARRETT CO.: Grantsville, Stone 14111 
(PENN). KENT CO.: Chestertown, Vanatta (CHRB). PRINCE GEORGE 
co.: Suitland, Meyer 9178 (ISC, WVA). QUEEN ANNES CO.: Chester 
River, Thatcher 5 (MIN). ST. MARY'S CO.: Patuxent River, Grether 
8276 (MICH). TALBOT CO.: Easton, Earle 810 (PENN). WASHINGTON 
co.: Boonsboro, Tidestrom 724a (US). WICOMICO CO.: Sharptown, 
Tidestrom 12228 (MICH). WORCESTER CO.: Ferry Creek, Redmond 
225 (MICH). Massachusetts: BERKSHIRE CO.: Williamstown, Ames 
(MICH). BRISTOL CO.: New Bedford, Bartlett 319 (IND, MICH). 
ESSEX CO.: Newburyport, Mock (NHA). FRANKLIN CO.: Mt. Toby, 
Elwell (MASS). HAMPDEN CO.: Granville, Seymour 197 (MASS). 
HAMPSHIRE CO.: Amherst, Elwell (MASS). MIDDLESEX CO.: Concord, 
Mann (HNH). NORFOLK CO.: Dedham, Greenman 2371 (MIN). 
PLYMOUTH CO.: Plymouth, Morong 1184 (US). WORCESTER CO.: 
Brookfield, Markert (MASS). Mississippi: CLARKE CO.: Burstone 
Cuesta, Jones 12050 (VDB). FORREST CO.: McCallum, Ray, Jr. 6045 
(MISSA). GEORGE CO.: Cedar Creek, Demaree 35222 (VDB). HARRISON 
co.: Gulfport, Bomhard 33 (NO). HINDS CO.: Natchez Trace Park- 
way, Price (MISSA). JACKSON CO.: Van Cleave, Demaree 82454 
(DUKE). JASPER CO.: Rose Hill, Reynolds 96 (MISSA). LAUDERDALE 
co.: Meridian, Hayes 20 (MISSA). LINCOLN CO.: Boque Chitto, Ray, 
Jr. 7830a (MISSA). PEARL RIVER Co.: Picayune, Ray, Jr. 6519 
(MISSA). STONE CO.: Perkinson, Ray, Jr. 7713 (MISSA). TISHOMINGO 
co.: Fisherman Ford, Ray, Jr. 7507 (MISSA, VDB). WAYNE CO.: Clara, 
Graham 118 (MISSA). New Hampshire: BELKNAP CO.: Center Harbor, 
Hodgdon, et. al. 9507 (NHA). CARROLL CO.: Albany, Pease 25358 
(MAINE). CHESHIRE CO.: Fitzwilliam, Hodgdon 7069 (HNA). GRAFTON 
co.: Hanover, Drew (HNH). HILLSBORO Co.: Manchester, Hodgdon 
& Kent 7070 (NHA). MERRIMACK CO.: Hooksett, Hall (NHA). ROCK- 
INGHAM CO.: Windham Center, Harris 430 (NHA). STRAFFORD CO.: 
Farmington, Lord (NHA). New Jersey: ATLANTIC CO.: Alsion, Crow 
126 (MSC). BERGEN CO.: Palisades, Gershoy (CU). BURLINGTON CO.: 
Pemberton, Palmer 89752 (MICH, MIN). CAMDEN CO.: Gibbsboro, 
Hynes (PENN). CAPE MAY CO.: Dennisville, Fender 5049 (PENN). 
CUMBERLAND CO.: Gouldtown, Long 575 (ARIZ, CAN, CU, IND, ISC, KY, 
NHA, NO, TENN, UARK, WVA). ESSEX CO.: Millburn, Rusby 1710 
co.: Fairmount, Good (CHRB). MIDDLESEX CO.: Spotswood, Laugh- 

356 Rhodora [Vol. 76 

ridge & Chrysler (CHRB). MONMOUTH CO.: Clarksburg, Fogg, Jr. 
14129 (PENN). MORRIS CO.: Longwood, Davidson 764 (CHRB). OCEAN 
C0.: South Lakewood, MacKenzie 3116 (US). PASSAIC co.: Utter- 
town Bog, Montgomery (CHRB). SALEM CO.: Auburn, Fogg, Jr. 6606 
(CHRB, PENN). SOMERSET CO.: Willowood estate, Blackburn (CHRB). 
SUSSEX CO.: Kittatinny Ridge, Davidson 1509 (CHRB). WARREN CO.: 
Sand Pond, Large & Clausen 1280 (CU). New York: BRONX CO.: 
Bronxdale, Burnham 428 (CU). CATTARAUGUS CO.: Rock City Park, 
Muenscher 15841 (CU). CHEMUNG CO.: Susquehana River, Lucy 6616 
(MU). COLUMBIA CO.: Ghent, McVaugh 881 (PENN). DELAWARE CO.: 
Deposit, Muenscher 16244 (cu). DUTCHESS CO.: Clove, Standley & 
Bollman 12138 (US). GREENE CO.: Winter Clove Mt., Gershoy (CU). 
NASSAU CO.: Woodbury, Stewart (CU). ONEIDA CO.: New London, 
House 27859 (CU). RICHMOND CoO.:. Haverstraw, Fosberg & Fosberg 
14897 (PENN). ROCKLAND CO.: Ramapo, Gershoy (CU). SCHUYLER CO.: 
Cayuta Lake, Muenscher & Brown 21703 (cU). STEUBEN CO.: Painted 
Post, Muenscher 15842 (CU). SUFFOLK CO.: Southampton, St. John 
2852 (CU). SULLIVAN CO0.: Camp Ranachqua, Weber 272 (isc). 
TIOGA CO.: Spencer, Hames & McDaniels 4765 (CU). TOMPKINS 
co.: Enfield, Palmer 928 (IND). ULSTER C0.: Lake Minnewaska, 
Gillis 3090 (MSC). WESTCHESTER CO.: Ossining, Gershoy (CU, MU). 

North Carolina: ALEXANDER CO.: Keever 21 (DUKE). BUNCOMBE 
co.: Asheville, McCarthy (US). BURKE CO.: Shortoff Mt., Wilbur 
7004 (VDB). CALDWELL CO.: Granite Falls, Randolph & Randolph 
1085 (CU). CHATHAM CO.: Costello 14 (DUKE). CLAY CO.: Perry 
Gap Road, Wilbur 1275 (MIN, DUKE). CUMBERLAND CO.: Rockfish 
Creek, Correll & Rankin 9053 (DUKE). DURHAM CO.: Catsburg, 
Wilbur 12230 (DUKE). EDGECOMBE CO.: Tar River, Fox & Whitford 
1746 (DUKE). FORSYTH CO.: Schallert 1092 (DUKE). GASTON CO.: 
Kings Mt., Oosting 1646 (DUKE). GRAHAM CO.: Thunderhind Mt., 
Beardslee & Kofoid (MU). GRANVILLE CO.: Tar River, Correll 516 
(DUKE, MASS). HARNETT CO.: Bunnlevel, Laing 1104 (MIN). HAY- 
woop co.: Waynesville, Standley 5401 (US). HENDERSON CO.: Hen- 
dersonville, Cottam 11140 (UT). JACKSON CO.: Wesner Bald, Ramseur 
171 (MAINE). JOHNSTON CO.: Neuse River, Fox & Whitford 1293 
(DUKE, WS). LEE CO.: Deep River, Beard 405 (KY). MCDOWELL CO.: 
Rogers 8358 (DUKE, US). MACON CO.: Mt. Satula, Harbison 1848 
(TENN). MADISON C0.: Hot Springs, Wehmeyer 533 (MICH). MIT- 
CHELL CO.: Roan Mt., Cannon 143 (CU). MOORE CO.: Carthage, 
Harriot (NO). ORANGE Co.: Hillsboro, Correll 325 (DUKE). POLK 
co.: Columbus, Townsend 236 (CU). ROCKINGHAM. CO.: Smith River, 
Correll & McDowell 10695 (DUKE). SCOTLAND CO.: Laurel Hill, 
Wiegand & Manning 2369 (cu). STANLY CO.: Badin, Fox, Godfrey 
& Boyce 1427 (DUKE, TENN, US, WS). STOKES CO.: Sauratown Mt., 
Radford 34665 (1sc). SURRY co.: Pilot Mt., Williams 296 (DUKE). 
SWAIN CO.: Andrews Bald, Bruhn 4 (TENN). TRANSYLVANIA CO.: 

1974] Genus Kalmia — Ebinger 357 

Rosman, Morley 891 (18C). UNION CO.: Rocky River, Ahles & Haesloop 
31521 (UARK). WAKE CO0.: Raleigh, Godfrey 3765 (US). WATAUGA 
co.: Boone, Ahles & Duke 43728 (PAC). WAYNE CO.: Mt. Olive, 
Rose 36 (MASS). WILKES CO.: Wyatt 832 (ARIZ, BRY, NHA, VDB). 
Ohio: ADAMS CO.: Mineral Springs, Herrick (0S). ATHENS CO.: 
Athens, Kellerman (0S). COLUMBIANA CO.: Sanders (OS). COSHOCTON 
co.: New Castle Twp., Selby (os). CUYAHOGA Co.: Bayville, Biebel 
(OS). FAIRFIELD C0.: Sugar Grove, Potzger 2969 (ND). GALLIA CO.: 
Jones (0S). GUERNSEY CO.: Liberty Twp., Thomas (0S). HOCKING 
co.: Clear Creek, Stuckey 2912 (0s). HOLMES CO.: Killbuck Twp., 
Herrick (08S). JACKSON CO.: Liberty Twp., Bartley & Pontius (os). 
JEFFERSON C0.: Amsterdam, Cusick (08). KNOX co.: Pipesville, 
Moldenke 12186 (0S). LAWRENCE CO.: Hanging Rock, Werner (os). 
LICKING C0.: Blackland Gorge, Stuckey 6603 (os). MEIGS CO.: 
Columbia Twp., Jones (0S). MONROE CO.: Adams Twp., Thomas 
(OS). MORGAN CO.: Bur Oak St. Park, ‘Herrich (0S). MUSKINGUM 
co.: Laurel Hill, Reed (0s). PERRY CO.: Jonathan Creek, Hicks & 
Dambach (os). PIKE CO.: Chimney Rocks, Crowl (0S). ROSS CO.: 
Tar Hollow, Crowl (os). SCIOTA CO.: Cooper Head Fire Tower, 
Demaree 10664 (os). TUSCARAWAS CO.: Wayne Twp., Brown (os). 
VINTON C0.: Lake Hope St. Park, Stuckey 6407 (0S). WASHINGTON 
co.: Vincent, Walpole 1129 (BLH). Pennsylvania: ADAMS CO: Birch 
River, Tanger 4434 (PENN, WS). ALLEGHENY C0.: Tom's Run Rd., 
Henry & Buker (PENN). ARMSTRONG CO.: Sherrett, Henry (PENN). 
BEDFORD CO.: Tatesville, Berkheimer 982 (PENN). BERKS CO.: Bech- 
telsville, Fender 1022 (PENN). BLAIR CO.: Blue Knob, Westerfeld 
16177 (PAC). BRADFORD Co.: Canton, Westerfeld 907 (PAC, PENN). 
BUCKS CO.: Janney, Long 76067 (PENN). BUTLER CO.: Clintonville, 
Brubaker 2537 (PENN). CAMBRIA CO.: Johnstown, Tinan (os). 
CAMERON CO.: Sinnemahoning Creek. Fogg, Jr. 11555 (PENN). 
CARBON CO.: Little Gap, Fogg, Jr. 11900 (MIN, PENN). CENTRE CO.: 
Yarnell, Keener 2581 (PAC). CHESTER CO.: Harmonvville, Fender 
1401 (PENN). CLARION CO.: Callensbure, Wood. Jr. 2170 (PENN). 
CLEARFIELD CO.: Troutville, Ehrle 2475 (PAC). CLINTON CO.: Boone- 
ville, Wahl 3198 (Pac). COLUMBIA co.: Mill Grove, Pohl 2554 
(PENN). CUMBERLAND CO.: Shipnensbere, Saxe, Jr. 36 (PENN). 
DAUPHIN Co.: Pillow. Berkheimer 12883 (PAC, PENN). DELAWARE 
co: Darby, Baker (Pac). ELK CO.: Medix Run, McClelland (PENN). 
FAYETTE CO.: Wvymps Gan, Bwinger (PENN). FOREST CO.: Brooks- 
ton. Shields D-2157 (PAC). FRANKLIN CO.: Mercersburg, Kriner 104 
(PAC). FULTON CO.: Deneen Gap, Westerfeld 7798 (Pac. PENN). 
GREENE CO.: Jefferson. Wherry & Bell (PENN). HUNTINGDON CO.: 
Maddensville, Westerfeld 9823 (PAC). JEFFERSON CO.: Sieel, Wahl 
10944 (PAC). JUNIATA CO.: Cross Keys, Westerfeld 14535 (PAC). 
LACKAWANNA CO.: Montdale, Adams 3542 (PENN). LANCASTER CO.: 
Collins, Brubaker 1521 (PENN). LAWRENCE CO.: McConnells Mills, 

358 Rhodora [Vol. 76 

Russell 1582 (PENN), LEBANON CO.: Cold Spring, Travis 1170 
(PENN). LUZERNE CO.: Mountaintop, Adams 3484 (PAC), LYCOMING 
co.: Cedar Run, Wahl 19261 (PAC). MIFFLIN CO.: Standing Stone 
Creek, Westerfeld 3961 (PAC). MONROE CO.: Scotrun, Niering 1214 
co.: Wind Gap, Schaeffer, Jr. 18252 (PENN). NORTHUMBERLAND C0.: 
Sunbury, Wagner 8987 (PENN). PERRY CO.: Bloomfield, Adams & 
Thebes 1232 (PENN). PIKE CO.: Porter’s Lake, DePue 136 (PAC). 
SCHUYLKILL C0.: Delano, Wagner 3227 (PENN). SNYDER CO.: Paxton- 
ville, Wade 783 (PENN). SOMERSET CO.: Jennerstown, Fogg, Jr. 15258 
(PENN). SULLIVAN CO.: Laporte, Westerfeld 381 (PAC, PENN). 
SUSQUEHANNA CO.: North Knob, Adams 3612 (PENN). TIOGA CO.: 
Mainesburg, Fender 1537 (PENN). UNION CO.: Laurelton, Ludwig 
1521 (PAC). VENANGO CO.: Franklin, Wahl 12633 (PENN). WARREN 
co.: Irvington, Moldenke 17479 (ND). WAYNE CO.: White Mills, 
Adams 4624 (PENN). WESTMORELAND CO.: Ligonier, Matthews 2399 
(PAC). WYOMING CO.: Crane Swamp, Stone (PENN). YORK CO.: 
Dillsburg, Hoover 3078 (PAC). 

Rhode Island: NEWPORT CO.: Tiverton, Mearns 211 (US). PROVI- 
DENCE CO.: Pascoag, Myers 350 (WVA). WASHINGTON CO.: Kingston 
Hill, Reynolds 649 (HNH). South Carolina: ANDERSON CO.: Carpen- 
ter’s Mill, Davis (MICH, MIN, UT). GREENVILLE CO.: Saluda River, 
Freckman 1239 (ISC). LEXINGTON CO.: Batesburg, McGregor 297 
(US). MARLBORO CO.: Little Pee Dee River, Radford 12494 (TENN). 
OCONEE CO.: Whitewater, Davis 392 (MASS). PICKENS CO.: Table 
Rock, Rodgers 540 (DUKE). SPARTANBURG CO.: Campobello, Erdman 
973 (BRY). SUMTER CO.: Poinsette St. Park, Holdaway (DUKE). 
Tennessee: ANDERSON CO.: Coal Creek, Kearney (MIN, OS, US). 
BLOUT CO.: Cades Cove, Wallace & Jennison 1621 (TENN). CAMP- 
BELL CO.: Jellico, Lamb & Williams (TENN). CHEATHAM CO.: Mar- 
rowbone Creek, De Selm 1258 (TENN). CLAIBORNE CO.: Lone Mt. 
Ridge, Kelley 2459 (TENN). COFFEE CO.: Old Stone Fort, Blum 3732 
(VDB). CUMBERLAND CO.: Ozone, Demaree 47496 (KY, NO, VDB). 
DAVIDSON CO.: Nashville, Hubbard 9112 (IND). DE KALB CO.: Smith- 
ville, De Selme 173 (TENN). DICKSON CO.: Sycamore School, Jobe 
(TENN). FENTRESS CO.: Clear Fork, Sharp 25648 (TENN). FRANKLIN 
co.: Elk River, Sharp & Adams 10785 (TENN). GRAINGER CO.: Blaine, 
Hodges 20 (TENN). GREENE CO.: Horse Creek Rd., Sherman & Sharp 
22280 (TENN). GRUNDY CO.: Mont Eagle, Stifler (CU). HAMILTON 
co.: Lookout Mt., Scribner (US). HARDIN CO.: Kral 36161 (VDB). 
HAWKINS CO.: Clinch Mt., Wolfe 19180 (TENN). HOUSTON CO.: Erin, 
Palmer 17610 (vs). JOHNSON CO.: Barclay 12B (TENN). KNOX CO.: 
New Hopewell, Sharp 40454 (TENN). LAUDERDALE CO.: Summertown, 
Postal (IND). LAWRENCE C0.: Shoal Creek, Sharp, Adams & Felix 
11058 (TENN). LEWIS CO.: Trace Creek, Sharp, Adams & Felix 
10065 (TENN). MARION CO.: Fosters Falls, Webb 368 (TENN). 

1974] Genus Kalmia — Ebinger 359 

MONROE CO.: Skona River, Galyon & Sharp 21603 (TENN). MORGAN 
CO.: University Forest, Shanks 3141 (TENN). POLK CO.: Ocoee River, 
Wilson 3726 (TENN). PUTNAM CO.: Monterey, Norris & Shanks 2475 
(TENN). RHEA CO.: Spring City, James (08). ROANE C0.: Caney Creek, 
Shanks 22078 (TENN). ROBERTSON CO0.: Pleasant View, Shanks, 
Woods, & Cooley 14776 (TENN). SEVIER CO.: Pittman Center, Jen- 
nison 3350 (TENN). UNICOI CO.: Rich Mt., Price 658 (DUKE). UNION 
co.: Norris Lake, Kelley (TENN). VAN BUREN CO.: Spencer, Shanks 
1288 (TENN). WASHINGTON CO.: Cherokee Mt., Sharp 3819 (TENN). 
WAYNE CO.: Butler Creek, Shanks, Woods & Cooley 9156 (TENN). 
WHITE CO0.: Caney Fork, Channell 8956 (VDB). WILLIAMSON CO.: 
Ferndale, Major 6 (vpB). Vermont: BENNINGTON C0.: Pownal, 
Eggleston (HNH, US). Virginia: ALBEMARLE CO.: Miller School, 
Tinsley (NMC). AUGUSTA CO.: Stuarts Draft, Churchill (Msc). 
BLANO CO.: Suiter, Kral 10453 (VDB). CRAIG CO.: Newcastle, Fogg, 
Jr. 13211 (PENN). FAIRFAX CO.: Great Falls, Bartlet 1783 (IND). 
GILES CO.: Salt Pond Mt., Fogg, Jr. 12688 (PENN). GREENE CO.: 
Bear Fence Mt., Fosberg 17205 (PENN). ISLE OF WIGHT CO.: Smith- 
field. Wise (KY). JAMES CITY C0.: Gordon Creek, Fernald & Long 
11393 (DUKE). KING GEORGE CO.: King George, Muller 3787 (ND). 
LOUDOUN CO.: Ashburn, Pratt (MU). MADISON CO.: Old Ray Mt., 
Fosberg 16909 (PENN). MONTGOMERY CO.: Blacksburg, Musselman 
645 (UWM). NANSEMOND CO.: Nansemond River, Fernald & Long 
13719 (US). NELSON CO.: Hall 3755 (VDB). NORFOLK CO.: Great 
Dismal Swamn. Fernald & Long 12147 (TENN). PITTSYLVANLA CO.: 
Smith Mt., Zobel (DUKE). ROANOKE CO.: Surarloaf Mt., Ebinger 4041 
(MU). ROCKBRIDGE CO.: G. Washington Nat’l. Forest, Hall 3716 
(VDB). ROCKINGHAM CO.: Shenandow Forest, Eggleston 18624 (us). 
RUSSELL CO.: Clinch Mt., Kral 23850 (VDB). SMYTH CO.: Pond Mt., 
Small (msc, US). WISE CO.: Norton, Pennell 11813 (Isc). West 
Virginia: BARBOUR CO.: Nestorville, Core 5922 (WVA). BERKELEY 
co.: Back Creek. Core 5847 (WVA). BRAXTON CO.: Little Birch, 
Harris (WVA). CALHOUN CO.: Pink, Harris (WVA). CLAY CO.: Clay, 
Core 6372 (WVA). DODDRIDGE CO.: West Union, Bartholomew (WVA). 
GRANT CO.: Dolly Sods Mt., Davis & Davis 7129 (WS). GREENBRIER 
co.: Muddy Creek Mt., Fox (WVA). HAMPSHIRE CO.: Ice Mt., Frye 
1269 (WVA). HARDY CO.: North Mt. Core & Ludwia 399 (WVA). 
HARRISON CO.: Duck Creek, Martin 327 (WVA). JEFFERSON CO.: 
Charlestown, Harper (CU). KANAWHA CO.: Nitro. Greenlee (WVA). 
MCDOWELL CO.: Panther, Music (WVA). MARION CO.: Pinesville, 
Bodola (WVA). MARSHALL CO.: Calis, Bartholomew 1941-217 (WVA). 
MERCER CO.: Pinnacle Rock, Fox (WVA). MINERAL CO.: Patterson 
Creek, Brown (WVA). MINGO CO.: Chattaroy, Bowen (WVA). 
MONONGALIA CO.: Cassville, Watkins (WVA). MONROE CO.: Wayside, 
Fox (WVA). MORGAN CO.: Langent, Core 5739 (WVA). NICHOLAS CO.: 
Calvin, Creasy (WVA). PENDLETON CO: Seneca Rocks, Clarkson 2001 

360 Rhodora [Vol. 76 

(WVA). POCAHONTAS CO.: Droop Mt., Ludwig 77 (WVA). PRESTON 
co.: Pisqah Rd., Meyer 247 (HNH, PENN, TENN, WVA). RANDOLPH 
co.: Roaring Plains, Core 3171 (WVA). RITCHIE CO.: Cairo, Good- 
win 251 (WVA). SUMMERS C0.: Barger Springs, Boone (WVA). 
TAYLOR CO.: Wickwire Run, Core 5475 (WVA). TUCKER CO.: Black- 
bird Knob, Gibson 1039 (WVA). UPSHUR C0.: Bucklin, Pollock (MIN, 
US). WAYNE CO.: Hurricane Creek, Lycan 59 (WVA). WEBSTER CO.: 
Camp Caesar, Fox (WVA). WETZEL CO.: Smithfield, Wagner (wva). 
WIRT CO.: Standingstone Creek, Bartholomew W-5007 (WvA). 

Mountain laurel commonly forms dense thickets in rocky 
and sandy forests throughout most of its range, particularly 
where there are openings in the canopy. It is also found in 
pastures and open fields and commonly forms thickets at 
the edge of roads. A detailed map of the known range of 
Kalmia latifolia is given by Kurmes (1967) who considers 
the report of this species from Ontario and New Brunswick 
to be an error, as is the report from the valley of the Red 
River in Arkansas (Britton, 1908). The report of this spe- 
cies from Canada probably originates from Pursh (1814), 
and it has been perpetuated by Brunet (1867), Sargent 
(1893), and more recently by Munns (1938). There is no 
evidence that this species is native to any part of Canada. 
It is possible that these early reports were based on culti- 
vated plants or large-leaved specimens of K. augustifolia 
(Macoun, 1884). Presently the species is known from south- 
ern and southeastern Maine (Fernald, 1911; Lamson- 
Scribner, 1891; Norton, 1913, 1935; and Perkins, 1935), 
southern New Hampshire (Hodgdon, 1955) and southern 
Vermont (Deane, 1899). 

The ecological requirements of Kalmia latifolia in the 
deciduous forests of eastern United States have been exam- 
ined by a few authors. Kurmes (1961) studied the ecology 
of mountain laurel in southern New England and deter- 
mined its exact range from herbarium material and field 
observations. The relationship of this species to water 
run-off and the water-holding capacity of the soil was 
studied by Colvin and Eisenmenger (1943) and Johnson 
and Kovner (1956). Studies have also been made of the 
rate of water-vapor loss from its leaves (Meyer, 1927), the 

1974] Genus Kalmia — Ebinger 361 

seasonal variation in the moisture content of its leaves 
(Reifsnyder, 1961) and seasonal changes in cold resistance 
(Parker, 1961). The competition of mountain laurel with 
tree seedlings and sprouts has also been studied. Wahlen- 
berg and Doolittle (1950) found that under normal condi- 
tions in the southern Appalachian mountains K. latifolia 
forms dense thickets that preclude any natural regeneration 
of timber trees. Their studies indicate that partial clearing 
of these thickets followed by planting of desirable tree spe- 
cies is an effective method of forest regeneration. They also 
found that mountain laurel has a mean annual height 
growth of 0.4 feet and a lateral growth of 0.3 feet. 

Kalmia latifolia is usually a tall, spreading shrub that 
throughout most of its range rarely exceeds a height of 
12 feet. In the fertile valleys between the Blue Ridge and 
the Alleghany Mountains, however, this species may attain 
the size of a small tree. Engelmann, Canby, and Gray 
(1877) observed a number of large individuals growing 
in the bottom of a dell, back of Caesar’s Head, on the ex- 
treme western border of South Carolina. One of the trunks 
measured 4 feet 11/4, inches in circumference about 1 foot 
above the ground while another in the same area was 3 feet 
4 inches above the first fork, and at 1 foot above the ground 
was 4 feet 4 inches in circumference. Later, Sargent (1893) 
records its maximum size as 30-40 feet tall with a diameter 
of 18-20 inches. The largest specimen on record, as listed 
by the American Forestry Association (1955), is a plant 
from the Great Smoky Mountains National Park with a 
circumference of 3 feet 6 inches, a height of 25 feet, and a 
spread of 45 feet. This specimen is probably located at 
Forney Ridge, Swain County, Tennessee (Barksdale & Jen- 
nison 1285). 

Besides the numerous studies concerned with the poison- 
ous properties of mountain laurel, some early attempts were 
undertaken to determine what other substances were pres- 
ent. These early studies reported the presence of arbutin, 
gums, and tannins (Kennedy, 1875), bark tannins (De- 
Graffe, 1896) and fat deposits (Stanley, 1931). More re- 

962 Rhodora [Vol. 76 

cently the effects of herbicides on Kalmia latifolia have 
been studied. Egler (1947, 1948) found that 2,4-D has no 
permanent effect, causing only temporary stunting and cur- 
vature of the foliage. Later, Egler (1949, 1950) found that 
a weak solution (.25%) of a 2,4-D and 2,4,5-T combination 
causes temporary cessation of growth, but a strong solution 
(596) is necessary to ensure death. 

The synonomy of mountain laurel is not very extensive. 
Most of the names proposed are for forms and varieties 
which are the result of the extreme variation that exists in 
flower color, leaf shape and size, plant size and pubescence. 
In 1940, Fernald described a variety (laevipes) of Kalmia 
latifolia from the coastal plain on southeastern United 
States. He considered this variety in its extreme develop- 
ment as having pedicels that are “quite smooth or with the 
merest suggestion of sessile glandular spots". In typical K. 
latifolia, in contrast, the pedicels are densely floccose and 
viscid with stalked glandular hairs. Many individuals with 
few or no glands on the pedicels are found in Indiana and 
Ohio, as well as on the coastal plain from Virginia to 
Louisiana. In some populations specimens with both gland- 
less and glandular pedicels are found. Also, Radford, Ahles, 
and Bell (1968) mention that this condition is very variable, 
with a tendency for pedicels to be floccose on plants in the 
mountains and less glandular on the coastal plain. Because 
of this variation it appears best to consider these individuals 
as an extreme that occurs in the southern and western parts 
of the range of K. latifolia. 

A total of 7 forms of Kalmia latifolia have been recog- 
nized by Rehder (1910, 1945, 1949). Many of these were 
originally described as varieties, but that classification is 
not appropriate for these taxa in either the horticultural or 
botanical sense. It is possible that these taxa could be 
treated as cultivars as done by Holmes (1956) and Dudley 
(1967), but most are better treated as forms as defined by 
Davis and Heywood (1963). The genetic evidence indicates 
that most of these taxa are not unique clones, but rather 
are sporadic variants distinguished by a single or a few 

1974] Genus Kalmia — Ebinger 363 

linked characters, without a distinct distribution. Also, 
they do not represent the normal extremes of most popula- 
tions, but are the result of the repeated association of re- 
cessive genes that are of a limited occurrence in some popu- 
lations. Of the 7 forms recognized by Rehder, 5 fit the 
criteria of a form as described above. The remainder, both 
variations in flower color (alba and rubra), unlike the 
forms, are not aberrant or disjunct, but are connected to 
the typical light pink flower by intermediate color types. 

Kalmia latifolia f. angustata Rehd. Jour. Arnold Arb. 26: 
481. 1945. TYPE: New Jersey: CAPE MAY CO.: Dennis town- 
ship, H. A. Scribner (GH). 

Kalmia latifolia B. salicifolia Forbes, Hort. Woburn. 93. 
1833. (nom. nud.). 

This very rare foliage form has narrowly oblanceolate to 
linear leaves that are 4-8 cm long and 4-10 mm wide. Liv- 
ing material of the type is cultivated at the Arnold Arbore- 
tum (Rehder, 1945). The only specimens of f. angustata 
seen by the author are cultivated plants. 

Common name: Willow-leaved Mountain Laurel. 

Kalmia latifolia f. fuscata (Rehd.) Rehd. Rhodora 12: 2. 

Kalmia latifolia var. fuscata Rehd. Moller's Deutsch. 
Gart. Zeit. 18: 578. 1903. TvPE: New York: Canaan Four 
Corners, Dr. Childs (GH). 

Kalmia latifolia var. coronata Gray, Amer. Nat. 2: 324. 
1868. TYPE: Massachusetts: woods near Framingham, 
James Parker (GH). 

Kalmia latifolia var. maculata Raf. Medical Flora 2: 17. 
1830. TvPE: unknown. 

This form is distinguished by white to pink flowers that 
have a heavily pigmented, usually continuous, brownish- 
purple or cinnamon band on the inside of the corolla at the 
level of the anther pockets. This band breaks up into 
brownish dots toward the base and the margin of the 
corolla. It also shows through to the outside of the bud 
giving it a muddy appearance. 

364 Rhodora [Vol. 76 

Some variation does exist in the size, shape and color of 
the band with an interrupted band existing in some of the 
specimens examined. According to Jaynes (personal com- 
munication) controlled crosses with specimens showing this 
banded characteristic strongly suggests that the character 
is determined by a single dominant gene. 

Common name: Banded Mountain Laurel, Crowned Moun- 
tain Laurel. 


Alabama: near Citronella, Milligan (US). Connecticut: near 
Chaplin, Jaynes (EIU). Massachusetts: near Framingham, Parker 
(GH); Wales, Clark (MASS); Guterman (MASS). New York: Brew- 
ster, Jaynes 240 (EIU). North Carolina: Highlands, Anderson (US). 
Ohio: Holden Arboretum, Mentor, Jaynes 94 (EIU). Pennsylvania: 
nursery in Stroudsburg, Jaynes 168 (EIU). 

Kalmia latifolia f. myrtifolia (Bosse) K. Koch, Dendrol- 
ogie 2(1) : 153. 1872. 

Kalmia latifolia var. myrtifolia Bosse, Vollst. Handb. 
Blumengart. 2: 348. 1841. Kalmia myrtifolia Andre, Revue 
Hort. 55: 10. 1883. TYPE: unknown, probably a living plant. 

Kalmia latifolia f. minor K. Koch, Dendrologie 2(1) : 153. 
1872. TYPE: unknown, probably a living plant. 

Kalmia latifolia var. nana Lavallée, Arbor. Segrez. 159. 
1877. (nom, nud.); Mouillefert, Traité Arb. Arbrisse 2: 
1027. 1897. TYPE: unknown, probably a living plant. 

This form is in all respects a miniature mountain laurel, 
being compact and slow-growing, rarely exceeding a height 
of 3 feet. In all specimens examined the leaves are much 
smaller than in typical Kalmia latifolia, averaging 2-4 cm 
long and about 1 cm wide. Also, flower size and length of 
the internodes are approximately 1/3 to 1/2 normal. Recent 
evidence (Jaynes, personal communication) suggests that 
this dwarf condition is controlled by a single recessive gene. 

According to Dudley (1967) this form has been grown 
since 1840 and is occasionally found in gardens where space 
is limited. Though occasionally mentioned in the early 
horticulture literature of Europe, the first reference to 
this form in North American literature appears to be by 

1974] Genus Kalmia — Ebinger 365 

Sprague (1871). The only report of wild plants of f. myrti- 

folia appeared in 1895 (Anonymus) which, according to the 

author, were identical to the cultivated dwarf form. 
Common name: Miniature Mountain Laurel, Dwarf 

Mountain Laurel. 


Connecticut: Portland, Jaynes 25 (EIU). Massachusetts: Arnold 
Arboretum, Hornby (MASS). 

Kalmia latifolia f. obtusata (Rehd.) Rehd. Rhodora 12: 2. 

Kalmia latifolia var. obtusata Rehd. Móller's Deutsch. 
Gart. Zeit. 18: 577. 1903. TYPE: Connecticut: near Pom- 
fret, Bowditch. 

Kalmia latifolia var. ovatifolia Raf. Medical Flora 2: 17. 
1830. (nom. nud.). 

This foliage form has leaves that are oval to oblong- 
obovate and rounded at both ends. In the specimens exam- 
ined the leaves average 4-7 cm long and 2.5-4 cm broad, are 
apiculate, and have short petioles that rarely exceed 1 cm 
in length. According to Rehder (1910) this form is slow 
growing, forms a compact bush, and has been cultivated at 
the Arnold Arboretum since 1886 when it was propagated 
from the type plant. Dudley (1967) suggested that the 
cultivar ‘Ovata’ which is available from some European 
sources is a synonym of this variant. 

Common name: Hedge Mountain Laurel. 

Connecticut: Riverton, Jaynes 291 (EIU). Massachusetts: 1 mi. 

east of Ashby, Ahles 65185 (MASS); Arnold Arboretum, Wyman 
2854 (MSC), Jaynes 238 (EIU). 

Kalmia latifolia f. polypetala (Nickolson) Beissner, 
Schelle, & Zabel, Handb. Laubholz-Benennung 386. 1908. 

Kalmia latifolia var. polypetala Nicholson, Hand-list 
Trees Shrubs Arb. 2: 49. 1896. Kalmia latifolia f. polype- 
tala (Nicholson) Rehd. Rhodora 12: 1. 1910. TYPE: Mas- 
sachusetts: near South Deerfield, Miss M. Bryant (GH). 

Kalmia latifolia var. monstruosa Mouillefert, Traité Arb. 

366 Rhodora [Vol. 76 

Arbrisse 2: 1027. 1897. TYPE: Unknown, probably a living 

This form has the corolla deeply divided into 5 narrow 
to fairly broad, strap-shaped petals. In some specimens 
these petals are extremely narrow and thread-like due to 
the revolution of the margins. Gray (1871) reports that 
these narrow petals resemble filaments and are sometimes 
tipped with imperfect anthers. More commonly the petals 
are broader, and a few specimens have been found in which 
the individual flowers appear much like apple blossoms. 
Other variations include an apetalous type which has been 
observed in some nursery stock from Massachusetts, and a 
cultivar named ‘Bettina’ (Dudley, 1967) in which the corolla 
is deeply lobed. According to Jaynes (personal communica- 
tion) the inheritance data fits the hypothesis that the genetic 
control of the polypetalous condition is by a single recessive 

This form was first described by Gray (1871) and later 
by Sargent (1890), while Bean (1897) mentioned that this 
unusual form flowers each year at the Kew Gardens. It was 
given the name var. polypetala by Nicholson (1896) and 
the name var. monstruosa by Mouillefert (1897), both 
based on Sargent's description of 1890. This taxon was 
first considered as a form by Beissner, Schelle, and Zabel 
(1903) who use the name polypetala and cite Sargent as 
the author. 

This form appears to occur most commonly in Massachu- 
setts. Stone (1909) reported it growing at the edge of a 
road in Leverett, not far from Mt. Toby, while Britton 
(1925) reports that Dr. Stone has found it from three dif- 
ferent stations, all on or near Mt. Toby. Mt. Toby is close 
to South Deerfield, the type locality of this form. 

Common name: Feather Petal Mountain Laurel. 

Massachusetts: Arnold Arboretum, Wyman 2458 (Msc); Leverett, 
Mt. Toby, Henry (MASS), Osmun (MASS), Stone (MASS), 67570 
(Mass). North Carolina: Danbury, Hanging Rock State Park, 
Rogers (EIU). 

1974] Genus Kalmia — Ebinger 367 

4. Kalmia angustifolia Linnaeus, Sp, Pl. 1: 391. 1753. 
Type: “Habitat in Pensylvania, Nova Caesarea, Nove- 
boraco" (LINN 560.2, lectotype, specimen on the right). 

Chamaedaphne angustifolia (L.) Kuntze, Rev. Gen. PLA 
388. 1891. 

Much branched shrub to 1.5 m tall; branchlets terete, 
reddish-brown, glabrous to pubescent and stipitate-glandu- 
lar, usually ascending; leaves in whorls of 3 (rarely a few 
alternate or opposite), slightly coriaceous, evergreen, flat, 
oblong to elliptic-lanceolate, 2-5 (rarely 7) cm long, 0.5-2 cm 
wide, usually lightly puberulent above with short white 
hairs, sometimes glabrate with age, midrib puberulent 
above, glabrous to puberulent beneath, sometimes stipitate- 
glandular; petioles to 1 (rarely 1.5) cm long, usually pu- 
berulent; inflorescence of numerous corymbiform racemes 
from the axils of last year’s leaves; pedicels 0.5-2 cm long, 
puberulent and stipitate-glandular, subtended by 2 bracte- 
oles in the axil of a lanceolate bract, 1-6 mm long; main 
axis of the inflorescence less than 1.5 cm long; calyx pu- 
berulent outside, glabrous within, usually green, sometimes 
the tip reddish, or reddish throughout, 3-6 mm broad, the 
lobes usually acuminate; corolla reddish-purple to pink 
(rarely white), puberulent outside, puberulent toward the 
base within, 6-13 mm broad; anthers purplish-brown, .5-1 
mm long; filaments puberulent on the lower half; capsule 
depressed globose, 3-5 mm broad, 2-3.5 mm long, puberu- 
lent and stipitate-glandular; seed stramineous, 0.6-1 mm 
long, and testa loose and extending past the ends of the 
seed. Chromosome number n — 12. 

Kalmia angustifolia is a common species in eastern North 
America. It is found from northern Ontario, Quebec and 
Labrador, through the Great Lakes region, and south along 
the east coast of the United States to the Carolinas. It is 
primarily a plant of bogs and other wet, open areas, but is 
also found in open woods and as a weed in open areas, some- 
times forming dense thickets. 

Genetically this species is most closely related to Kalmia 
latifolia. When K. angustifolia is the male parent the cross 

368 Rhodora [Vol. 76 

is not successful because the sheep laurel pollen will not 
grow down the style of mountain laurel (Jaynes, 19682). 
The reciprocal cross, however, resulted in about 18% of 
the flowers producing capsules which contained some viable 
seed. These resulting interspecific hybrids were intermedi- 
ate to the parental types, although leaf arrangement was 
primarily alternate, as in K. latifolia. Jaynes (1968a) also 
reported that K. angustifolia could be crossed with both 
K. hirsuta and K. polifolia. Both crosses produced mostly 
weak, yellow-green individuals that died within the year. 
A few individuals lived longer, and some produced flowers 
with sterile pollen. 

Small (1903, 1914) considered this taxon as two separate 
species, a procedure not followed by subsequent authors. 
Both Fernald (1950) and Gleason (1952) considered this 
complex as being one species with two fairly distinct vari- 
eties, a view that has recently been followed by Radford, 
Ahles, and Bell (1968). Considering the genetic and mor- 
phological similarities of these taxa it is more realistic to 
follow the latter view and consider it as a single species 
with two fairly distinct varieties. 


l. Calyx densely stipitate-glandular; mature leaves not 
puberulent beneath. ............ 4a. var. angustifolia. 
l. Calyx lacking stipitate-glands, or nearly so; mature 
leaves densely puberulent beneath. . 4b. var. caroliniana. 

4a. Kalmia angustifolia L. var. angustifolia. 
Kalmia angustifolia B carnea Ait. Hort. Kew. 2: 64. 1789. 
Kalmia angustifolia var. hirsuta Jager, Ziergeholze 278. 

1865. Kalmia angustifolia f. hirsuta Voss, Blumengart. 1: 
591. 1894. (without basonym). 

Kalmia angustifolia var. lucida Lavalée, Arbor. Segrez. 
159. 1877. (nom. nud.). Kalmia angustifolia f. lucida 
Beissner, Schelle, & Zabel, Handb. Laubholz-Benennung 386. 
1903. (nom. nud.). 

1974] Genus Kalmia — Ebinger 369 

Kalmia angustifolia var. minima Bosse, Vollst. Handb. 
Blumengart. 2: 347. 1841. 

Kalmia angustifolia var. nana Bosse, Vollst. Handb. 
Blumengart. 2: 347. 1841. Kalmia angustifolia f. nana 
Voss, Blumengart. 1: 591. 1894. (as snyonym). 

Kalmia angustifolia B ovata Pursh, Fl. Amer. Sept. 296. 

Kalmia angustifolia f. procumbens Beissner, Schelle, & 
Zabel, Handb. Laubholz-Benennung 386. 1903. (nom. 

Kalmia angustifolia, var. rosea Forbes, Hort. Woburn. 93. 
1833. Kalmia angustifolia f. rosea Beissner, Schelle, & 
Zabel. Handb. Laubholz-Benennung 386. 1903. (nom. 

Kalmia angustifolia var. variegata Forbes, Hort. Woburn. 
93. 1833. 

Kalmia elliptica Raf. Autikon Bot. 87. 1840. (as syno- 

Kalmia ferruginea Raf. Autikon Bot. 86. 1840. TYPE: 

Kalmia media K. Koch, Dendrologie II; 1: 153. 1872. 
Kalmia angustifolia f. media Beissner, Schelle, & Zabel, 
Handb. Laudholz-Benennung 386. 1903. (nom. nud.) TYPE: 

Kalmia intermedia Lange Ind. Sem. Hort. Hauniensi 
Nom. emendata 3. 1873. Kalmia glauca B intermedia Lange 
Ind. Sem. Hort. Hauniensi Nom. emendata 3. 1873. (as 
synonym). TYPE: a herbarium specimen in the Botanical 
Museum of Copenhagen (C) labeled Kalmia intermedia 
Lge ad int. Hort. bot. Haun. 16. Juni 1873, sem. ex hort. 
Paris s.n. 

Kalmia angustifolia B oleifolia Pers. Syn. Pl. 1: 4T". 
1805. Kalmia oleifolia (Pers.) Bosse, Vollst. Handb. Blu- 
mengart. 2: 347. 1841. (as synonym). TYPE: unknown. 

Kalmia angustifolia var. pumila Forbes, Hort. Woburn. 
93. 1833. Kalmia pumila Bosse, Vollst. Handb. Blumengart. 
2:347. 1841. (as synonym). 

370 Rhodora [Vol. 76 

Kalmia pumila rubra Bosse, Vollst. Handb. Blumengart. 
2:347. 1841. (as synonym). 

Kalmia pumila serotina Bosse, Vollst. Handb. Blumengart. 
2: 347. 1841. (as synonym). Kalmia angustifolia f. pumila 
Voss, Blumengart. 1: 591, 1894. (without basonym). 
TYPE: unknown. 

Kalmia angustifolia « rubra Ait. Hort. Kew 2: 64. 1789. 
Kalmia angustifolia var. rubra (Ait.) Loddiges, Bot. Cab. 
6: 502. 1821. Kalmia rubra (Loddiges) Bosse, Vollst. 
Handb. Blumengart. 2: 347. 1841. (as synonym). Kalmia 
angustifolia f. rubra Voss, Blumengart. 1: 591. 1894. 
(without basonym). TYPE: unknown. 

Kalmia angustifolia var. serotina Bosse, Vollst. Handb. 
Blumengart. 2: 347. 1841. Kalmia serotina Bosse, Vollst. 
Handb, Blumengart. 2: 347. 1841. (as synonym). TYPE: 

Mature leaves glabrous or with scattered stalked glandu- 
lar hairs beneath, usually not puberulent; calyx puberulent 
and strongly stipitate-glandular; corolla glabrous or lightly 
puberulent on the outside. 

Distribution: Bogs, Swamps, and other wet open areas 
from Ontario east to the Atlantic Ocean and southeast 
through the Great Lakes to Pennsylvania and south along 
the coast to Virginia. 

Common name: Sheep Laurel, Lambkill, Sheepkill, 
Wicky, Narrow-leaved Kalmia, Dwarf Laurel, Pig Laurel. 

UNITED STATES: Connecticut: FAIRFIELD CO.: Trumbull, Eames 
2809 (CONN). HARTFORD CO.: Kensington, Cowles (MU). LITCHFIELD 
CO.: Salisbury, Ebinger 697 (MU). NEW HAVEN co.: New Haven, 
Safford 163 (US). NEW LONDON CO.: Groton, Jausson (CONN, UT). 
TOLLAND CO.: Donovans Bog, Travis 1959 (PENN). WINDHAM CO.: 
Hasting’s Woods, Petersen (VDB). Delaware: NEW CASTLE CO.: Hog- 
swamp Road, Tatnall 4858 (GH). SUSSEX CO.: Canby (Us). Maine: 
ANDROSCOGGIN CO.: Lewiston, Bean 14793 (VDB). AROOSTOOK CO.: 
Knoles Corner, Neal 1907 (MAINE). CUMBERLAND CO.: Cape Eliza- 
beth, Gayle 766 (Us). HANCOCK CO.: Bar Harbor, Rossbach 2757 
(ACAD, NCU). KENNEBEC CO.: Clinton, Bean (MAINE). KNOX CO.: 

1974] Genus Kalmia — Ebinger 371 

Glencove, F'riesner 7721 (MU, RENO, UT, WS, WVA). LINCOLN CO.: 
Ocean Point, Fassett 3946 (DUKE). OXFORD CO.: White Mt. Nat. 
Forest, Richards 6066 (MAINE). PENOBSCOT CO.: Old Town, Davis 
(MAINE). PISCATAQUIS CO.: Sangerville, Fernald 127 (MIN). SAGA- 
DAHOC CO.: Five Islands, Rehder 1018 (GH). SOMERSET CO.: Baker 
Brook, St. John & Nichols 2431 (CAN, NY). WALDO CO.: Searsmont, 
Rossbach 1932 (ACAD, MAINE). WASHINGTON CO.: Joe Dyer's Point, 
Walker 3039 (MAINE). YORK CO.: Sand Pond, Richards 5951 (MAINE). 
Maryland: ANNE ARUNDEL C0.: Severn Run, Adams & Wherry 3200 
(GH). cECIL co.: North East, Long 54422 (GH). CHARLES CO.: 
Waldorf, Niewwland (ND). PRINCE GEORGES CO.: Airport Bog, Her- 
man 11546 (NY). Wicomico CO.: Sharptown, Tidestrom 12229 (GH). 
WORCESTER CO.: Furnace, Redmond 237 (MICH). Massachusetts: 
BARNSTABLE CO.: Falmouth, Greenman 3043 (MIN). BERKSHIRE CO.: 
Great Barrington, Eggleston 21413 (MIN, NY). ESSEX CO.: Marble- 
head, Nelson 341 (CAN). FRANKLIN CO.: Mt. Toby, Seymour 2566 
(DUKE, MASS). HAMPDEN CO.: Granville, Seymowr 137 (MASS, MIN, 
Townsend, Ahles 65188 (MASS). NANTUCKET CO.: Polpis, Mackeever 
504 (PENN). NORFOLK CO.: Medfield, Batchelder 234 (NCU). 
PLYMOUTH CO.: Ellisville, Erdman 4223 (BRY). WORCESTER CO.: 
Mount Wachusett, Palmer 43196 (MIN). Michigan: ALCONA CO.: 
Lincoln, Voss 4583 (MICH). ALPENA CO.: Alpena, McVaugh 
11405 (MICH). | ARENAC CO.: Omer, Voss 6794 (MICH). 
CHARLEVOIX CO.: Garden Isl, Cain 149 (TRT). CHEBOYGAN CO.: 
Wolverine, Gleason & Gleason, Jr. 248 (GH, IND, ISC, NY, WVA). 
CRAWFORD CO.: Grayling, Voss 3098 (MICH). 10sCO CO.: Sand Lake, 
Dreisbach 7243 (CONN). KALKASKA CO.: Zavitz 1168 (QUK). MIS- 
SAUKEE CO.: Moorestown, Voss 3087 (MICH, OS). MONTMORENCY CO.: 
Hagenah 4936 (BLH). oscopa Co.: Rhodes Lake, Voss 3578 (MICH). 
OTSEGO CO.: Hardwood Lake, MeVaugh 10888 (BLH, CAN, MICH, MO). 
PRESQUE ISLE CO.: Emma, Clover 26193 (MICH). ROSCOMMON CO.: 
Roscommon, Voss 4258 (MICH). SAGINAW CO.: Sage Lake, Case, Jr. 
(MICH). New Hampshire: BELKNAP CO.: Crystal Lake, Hodgdon 
& Steele 9124 (NHA). CARROLL CO.: Albany, Gale (NHA). CHESHIRE 
co.: Winchester Manning (BRY). coos Co.: Randolph, Pease 4696 
(GH, MICH). GRAFTON CO.: Pike, Mathias 163 (MIN). HILLSBORO 
co.: Mt. Monadnock, Andrews 895 (ORE). MERRIMACK CO.: Wilmot, 
Hodgdon & Steele 9949 (NHA). ROCKINGHAM CO.: Harris 224 (NHA). 
STRAFFORD CO.: Rochester, Barrett & Hodgdon 15522 (NHA). SULLI- 
VAN CO.: Springfield, Hodgdon & Steele 9129 (NHA). New Jersey: 
ATLANTIC CO.: Hammonton, Keefe 123 (UWM). BERGEN CO.: Carl- 
stadt, Niederer (NY). BURLINGTON CO.: Rancocas River, Leonard 
6314 (US). CAMDEN CO.: Atco, Painter 684 (MIN, US). CAPE MAY 
co.: Belleplain, Fender 4916 (PAC, PENN), CUMBERLAND CO.: Mill- 
ville, Adams 324 (GH, PENN). ESSEX CO.: Franklin, Rusby (CHRB). 

372 Rhodora [Vol. 76 

GLOUCESTER CO.: Repaupo, Long 16065 (GH). MIDDLESEX CO.: Mill- 
town, House (US). MONMOUTH CO.: Farmingdale, Long & Brown 
9557 (PENN). MORRIS CO.: Mt. Hope, Britton (CHRB). OCEAN CO.: 
Lakewood, Mackenzie 3112 (Us). PASSAIC co.: Uttertown Bog, 
Montgomery (CHRB). SALEM CO.: Auburn, Fogg, Jr. 6616 (PENN). 
SOMERSET CO.: Sourland Mts., Fogg, Jr. 12406 (PENN). SUSSEX CO.: 
Cranberry Lake, Mackenzie 717 (ARIZ, IND). 

New York: ALBANY CO.: Karmer, House 7850 (CAN). CHENANGO 
co.: Plymouth Pond, Wiegand 6976 (GH). CLINTON CO.: Rudd 816 
(US). COLUMBIA CO.: Ghent, McVaugh 362 (PENN). DUTCHESS CO.: 
Susan Court, Ahles 64464 (MASS). ESSEX CO.: Lake Harris, House 
8072 (CAN). FRANKLIN CO.: Upper Saranac Lake, Muenscher, et. al. 
476 (NY). FULTON CO.: Gloversville, Haught (WVA). GREENE CO.: 
Coville (GH). HAMILTON CO.: Little Tupper Lake, Fosberg 16101 
(PENN). JEFFERSON CO.: Perch Lake, Beschel 17144 (QUK). LEWIS 
co.: Martinsburg, ‘Hotchkiss 2654 (GH). NASSAU CO.: Massapequa 
St. Park, Cain 93 (MO, PENN, TENN). ONEIDA CO.: Sylvan Beach, 
Maxon (US). ORANGE CO.: Sutherland Pond, Mearns 125 (US). 
RENSSELAER CO.: West Sand Lake, Brown 652 (DUKE). RICHMOND 
co.: Staten Isl, Dowell & Davis 2224 (GH). ST. LAWRENCE CO.: 
Norfolk, Phelps 756 (GH, NY, US). SARATOGA CO.: Corinth, Whitney 
4124 (PENN). SUFFOLK CO.: South Haven, Moldenke 18685 (NY). 
SULLIVAN CO.: Wurtshore, Hess 1267 (ARIZ). ULSTER CO.: Highland, 
Chrysler (CHRB). WARREN CO.: Luzerne, Fogg, Jr. 20309 (PENN). 
WASHINGTON CO.: Lake Nebo, Burnham 2756 (MIN). Pennsylvania: 
BERKS C0.: Hamburg, Berkheimer 1734 (PENN). BRADFORD CO.: Troy, 
Beardslee (MU). BUCKS CO.: Bristol, Adams 4514 (MAINE, PENN). 
CARBON CO.: Little Gap, Adams 3331 (GH, ISC, PENN). CHESTER CO.: 
Warwick, Wherry (GH). COLUMBIA CO.: Centralia, Fogg, Jr. 14256 
(NY, PENN). CUMBERLAND CO.: Sand Springs, Ward (PAC), DELA- 
WARE CO.: Tinicum, Baker (PENN). FRANKLIN CO.: Caledonia St. 
Park, Overhots (PAC). LACKAWANNA CO.: West Mt., Glowenke 6528 
(MIN, NY). LANCASTER CO.: Smithville, Small (US). LEHIGH CO.: 
Allentown, Dowell 1115 (Us). LUZERNE CO.: Freeland, Orton (PAC). 
LYCOMING CO.: Bald Eagle Mt., Keener 2561 (PAC). MONROE CO.: 
Tannersville, Swartley 791 (MAINE). NORTHAMPTON CO.: Daniels- 
ville, Schaeffer, Jr. 6184 (PAC). NORTHUMBERLAND CO.: Montandon, 
Reed 235 (CHRB). PIKE CO.: Twin Lakes, De Pue 878 (PENN). 
SCHUYLKILL CO.: Tamaqua, Kriebel 8586 (ND). SOMERSET CO.: 
Salisbury, Caffrey (PAC). SULLIVAN CO.: Double Run, Fogg, Jr. 
14437 (PENN). WAYNE CO.: Big Spring Pond, Glowenke 2434 
(PENN). WYOMING CO.: Falls, Glowenke 2023 (PENN). YORK CO.: 
McCall’s Ferry, Heller & Halbach (MIN). Rhode Island: KENT CO.: 
Warwick, Congdon (08). NEWPORT CO.: Newport, Mearns 254 (GH). 
PROVIDENCE CO.: Slaterville, Palmer 43348 (ISC, MIN). WASHINGTON 
co.: South Kingstown, Collins (CONN). Vermont: ADDISON CO.: 

1974] Genus Kalmia — Ebinger 373 

Bristol Pond, Eggleston 21453 (US). BENNINGTON CO.: Manchester, 
Day 267 (GH). CALEDONIA CO.: Barnet, Blanchard (HNH, MO). 
CHITTENDEN CO.: S. Burlington Twp., Charette 2190 (BRY, MIN). 
FRANKLIN CO.: East Berkshire, Clausen (MO, US). ORANGE CO.: 
Fairlee, Grover (MU). RUTLAND CO.: Middletown Springs, Carpenter 
(NCU). WASHINGTON CO.: Roxbury, Countryman 820 (HNH, NHA). 
WINDHAM CO.: Westminster, Robinson 30 (GH). WINDSOR CO.: 
Reading, Kittredge (NY). Virginia: ISLE OF WIGHT Co.: Franklin, 
Heller 1124 (GH, MIN, MO, PENN, US, WS); Lee’s Mill, Fernald & 
Long 12146 (GH, MO, NY, US), Harvill 15946 (NCU); Zuni, Fernald, 
Griscom & Long 6665 (GH, NY), 6666 (GH, PENN). NANSEMOND CO.: 
Cherry Grove, Fernald & Long 11392 (DUKE, GH, US); County Rd. 
665, Duke 1067 (NCU); South Quay, Fernald & Long 10366 (GH, 

CANADA: Labrador: Assizes Isl., Potter & Brierly 3424 (GH); 
Cape St. Charles, Potter & Brierly 3427 (GH); Cartwright, Brown 
(GH); Forteau, Long 28832 (GH); Goose Bay, Gillett & Findley 5352 
(MIN, MO, TRT, WS), 5584 (ACAD, NY, US); Kenamu River, Gillett 
& McKay 5831 (ACAD, GH); St. Clary’s River, Wallace 23 (CONN). 
New Brunswick: ALBERTA CO.: Grindstone Isl. Christie 1575 (NBM). 
CARLETON CO.: East Wolf Isl, Hodgdon & Pike (NCU). CHARLOTTE 
co.: Deer Isl., Squires & Squires (NBM), GLOUCESTER CO.: Four 
Roads, Roberts & Pugh 65-4358 (CAN). KENT CO.: Richibucto, 
Scoggan 12135 (CAN). KINGS CO.: Havelock, Perry (ACAD). MADA- 
WASKA CO.: St. Leonard, Malte 294 (CAN). NORTHUMBERLAND CO.: 
Mt. Carleton, Smith & Clattenburg 20066 (ACAD, QUK). QUEENS CO.: 
Mill Brook, Squires & Christie (NBM). RESTIGOUCHE CO.: Dalhousie, 
Scoggan 11721 (ACAD, CAN). ST. JOHN CO.; Mispec, Crawford & 
Patterson (ACAD). SUNBURY CO.: Mill Settlement, Christie 245 (NBM). 
WESTMORLAND CO.: Johnson Lake, Beschel 5508 (QUK). Newfound- 
land: BONAVISTA NORTH DIST.: Glovertown, Smith, Squires, & Smith 
116 (ACAD), 135 (ACAD). BONAVISTA SOUTH DIST.: Lethbridge, 
Rouleau 8370 (NFLD). BURGEO-LA POILE DIST.: Osmond, Rouleau 7375 
(NFLD, US). CARBONEAR DIST.: Old Perlican, Torrey 25 (CAN, GH). 
FERRYLAND DIST.: Witless Bay, Rouleau 5364 (NFLD). FOGO DIST.: 
Stag Harbour, Ludlow 125 (ACAD). FORTUNE BAY-HERMITAGE DIST.: 
Wallace Cove, Rouleau 8089 (NFLD). GANDER DIST.: Gander, Rouleau 
6133 (CAN, NFLD, US). GRAND FALL DIST.: Overflow Pond, Rouleaw 
7961 (NFLD). HUMBER DIST.: Corner Brook, Rouleau 3960 (ACAD, 
CAN, GH, NFLD, NY, US). ST. BARBE DIST.: Bonne Bay, Bishop 471 
(GH, CAN). ST. JOHN’s DIST.: St. John's, Robinson & Schrenck 41 
(GH, CAN, MIN, MO, NY, P). TRINITY SOUTH DIST.: Whitbourne, 
Rouleaw 6069 (NFLD). TWILLINGATE DIST.: Loon Bay, Rouleau 4866 
(NFLD). WHITE BAY DIST.: St. Anthony, Savile & Vaillancourt 2561 
(NCU, NFLD, NY). Nova Scotia: ANNAPOLIS CO.: Belleisle, Bent 
12723 (ACAD). ANTIGONISH co.: Linwood, Smith, et. al. 10765 

374 Rhodora [Vol. 76 

(MAINE). CAPE BRETON CO.: Frenchvale, Smith, et. al. 15094 (ACAD). 
Wharton, Schofield 3197 (ACAD). DIGBY CO.: Corberrie, Jack 3121 
(MAINE, NY, US). GUYSBOROUGH CO.: Canso, Rousseau 35450 (CAN). 
HALIFAX C0.: Halifax, Jack 3235 (NY, TENN). HANTS CO.: Bishop- 
ville Rd., Wellwood 12662 (ACAD). INVERNESS CO.: North Mt., Smith, 
et. al. 11275 (ACAD). KING CO.: Wolfville, Anderson (ACAD). 
LUNENBURG C0.: Beck Lake, Smith, et. al. 18551 (ACAD, NCU). PICTOU 
co.: Westville, Sampson 28 (ACAD). QUEENS CO.: Tobeatic Lake, 
Martin 5210 (CAN). RICHMOND CO.: Grand Lake, Ershine 51993 
(QUK). SHELBURN C0.: Barrington Passage, Macoun 81371 (CAN). 
VICTORIA CO.: Cape Breton, Senn 1553 (MIN, US). YARMOUTH CO.: 
Seal Isl, Erskine 54.977 (ACAD). Ontario: BRUCE CO.: Big Bay, 
Krotkov 9295 (NY). CARLETON CO.: Wright’s Cove, Minshall 236 
(MIN, NY). COCHRANE CO.: Abitibi, Pritchard 208A (MASS). DUNDAS 
co.: Elma, House 23357 (PENN). FRONTENAC CO.: Kingston, Fowler 
(US). HALIBURTON CO.: Minden Twp., Roberts (TRT). KENORA DIST.: 
Attawapiskat River, Porsild, et. al. 19986 (CAN). LENNOX CO.: Skoota- 
matta, Jackson (TRT). MANITOULIN DIST.: Honora, Soper & Bell 9819 
(CAN, QUK, TRT). MUSKOKA DIST.: Sparrow Lake, Walden 110 (TRT). 
NIPISSING DIST.: Stuarts, Ross 2 (TRT). ONTARIO CO.: Cannington, 
Thompson 516 (TRT). PARRY SOUND DIST.: Ahmic Lake, Soper 5225 
(CAN, TRT). PRESCOTT CO.: Alfred Twp., Calder & Cody 922 (PENN). 
RENFREW CO.: Renfrew, Stroud (TRT). STORMONT CO.: Newington, 
Jenkins 8297 (NCU). SUDBURY DIST.: Sudbury, Ostenfeld 613 (CAN). 
THUNDER BAY DIST.: Longlac, Baldwin & Breitung 3693 (CAN, TRT). 
TIMISKAMING DIST.: Cattle Isl., Moss 77 (ALTA). WELLAND C0.: 
Niagara, Zenkert (OAC). Prince Edward Island: KING CO.: Dundee, 
Fernald, Long, & St. John 7899 (GH). QUEENS C0.: Brackley Beach, 
Grandtner 18518 (CAN). Quebec: ABITIBI CO.: Amos, Morton 11035 
(US). ARGENTEUIL CO.: Lachutem, Porsild 6337 (CAN). BERTHIER 
co.: Lanoraie, Scoggan 370 (CAN). BONAVENTURE CO.: New Rich- 
mond, Chrysler 1228 (CHRB). BROME CO.: Bolton, Knowlton (GH). 
CHICOUTIMI CO.: Lac Kenogami, Lavoie, Doyon, & Lord 600706-01 
(ACAD, TRT). GASPE WEST CO.: Mt. Albert, Cottam 11852 (UT). 
GATINEAU CO.: Wilson Corners, Mitchell 1750 (TENN). KAMOURASKA 
co.: Ouelle, Gauthier 65-207 (QUK). LABELLE CO.: Nomininque, Roy 
1053 (ACAD). LEVIS CO.: Chaudiere, Forbes (CAN). MAGDALEN ISLAND 
co.: Amherst Isl, Potter & Brierly 3425 (NMC, UT). MASKINONGE 
co.: Metis Beach, DuBoulay & DuBoulay 2589 (QUK). MATANE CO.: 
Lake Montagne, Boivin 786 (PENN). MATAPEDIA CO.: Saint Zenon, 
Blain 321 (CAN). MEGANTIC CO.: Black Lake, Blais, Deshaies, & 
Forest 10379 (CAN, QUK, SASK). MISSISQUOI CO.: Farnham, Maycock 
1863 (QUK). MONTCALM CO.: Lac Tapani, Pennell 16678 (NY). 
MONTMORENCY CO.: Boischatel, Cauchon, et. al. 69-111 (MU, TRT). 
PORTNEUF CO.: Marie-Victorin 9437 (WS). RICHELIEU CO.: Saint-ours, 

1974] Genus Kalmia — Ebinger 975 

Rolland-Germain 48518 (GH). RIMOUSKI CO.: Cap Enrage, Rousseau 
26877 (GH). RIVIERE DU LOUP CO.: Soper 3670 (TRT). SAGUENAY CO.: 
Mingan, Townsend (GH); Natashquan, Abbe 1188 (GH); Romaine, 
St. John 90652 (GH); Round Lake, Gillett 5208 (GH); Seven Isl., 
Harper 3309 (CAN, HNH, PENN, US), Robinson 769 (CAN, GH, NY). 
SHERBROOKE CO.: Orford, Martineau 35 (CAN). TEMISCOUATA CO.: 
Trois Pistoles, Fox (TRT). TERREBONNE CO.: St. Sophe, Wahl 19419 
(PAC). TIMISKAMING CO.: Ottawa River, Baldwin 5823 (CAN, SASK). 
TWO MOUNTAINS CO.: La Trappe, Louis-Marie 10 (GH). WOLFE CO.: 
Lake Nicolet, Brisson & Hamel] 11295 (ACAD, OAC, SASK, TRT). 

The synonomy of this taxon is fairly extensive. Numerous 
varietal and forma names have been proposed for the vari- 
ation that exists, and some of these names have been ele- 
vated to specifie rank. Many of these lack descriptions or 
are listed as synonyms under a variety or form name. Ex- 
cept for these subspecific categories only a few other names 
have been applied to this taxon. One of these was when 
Kuntze (1891) transferred the specific epithet angustifolia 
to another genus while the others were proposed by Rafin- 
esque (1840). 

Most of the subspecific categories that have been pro- 
posed for Kalmia angustifolia var. angustifolia are for 
variation and extremes in flower color (carnea, rosea, 
rubra), variation in size and habit of the plant (media, 
minima, nana, procumbens, pumila) and variation in shape 
and color of the leaves (lucida, oleifolia, ovata, variegata) . 
In general, the differences in flower color appears to repre- 
sent natural variation within populations and the above are 
better treated as cultivars than as forms. This is also true 
of the variation in leaf shape and color, and, in part, for 
variation in plant height. Holmes (1956) discusses some 
of these cultivars. 

The dwarf condition reported for Kalmia angustifolia 
var. angustifolia is either the result of selection of more 
northern or high altitude sources or because of inbreeding 
depression. Jaynes (19682) found that seedlings from 
selfed plants average about half the height of seedlings 
from outcrossed plants. He also found that seed set from 
self-pollinated plants is 85-90% less than that from cross 

376 Rhodora [Vol. 76 

pollination and that there is a marked reduction in survival 
of the seedlings from selfed plants. This indicates that this 
species is only weakly self compatible and that outcrossing 
usually occurs under natural conditions. 

Kalmia angustifolia var. angustifolia commonly occurs 
in bogs, swamps, and other wet places, usually forming 
dense thickets around ponds and lakes. It is also found in 
open woods, as a weed in fields, and in moist openings of 
pine savannas. It is distributed from the northeastern part 
of the Michigan peninsula and the eastern half of Ontario, 
east through Quebec, the Maritime Provinces, and New- 
foundland; north to at least the Attawapiskat River (Ken- 
ora District) in Ontario and to Goose Bay and Cartwright 
in Labrador; and south in eastern United States through 
New England and eastern New York, eastern Pennsylvania 
and Maryland to the coast in New Jersey and Delaware and 
the southeastern tip of Virginia. One specimen (Davis & 
Davis, 4691) has been seen which is reputed to have come 
from Randolph Co., West Virginia, but this collection should 
be considered doubtful. This species has also been reported 
from Indiana, but according to Deam (1940) this is an 

It is possible that extracts from sheep laurel may inhibit 
the growth of other plants, particularly coniferous trees, 
thereby maintaining open areas for extended periods of 
time. Peterson (1965) found that the leaves of this species 
contain a substance that hinders the primary root develop- 
ment of black spruce by destruction of the epidermal and 
cortical cells. He further suggests that this may be the 
reason for the abnormally poor growth of trees associated 
with Kalmia angustifolia on upland sites in the boreal 
region of eastern Canada. 

Studies by Jaynes (personal communication) reveal that 
some incompatibility exists between the two varieties of 
Kalmia angustifolia. This incompatibility is highly variable, 
depending upon the parents used, with the resulting off- 
spring varying from completely sterile to highly fertile. 
Many of these hybrids produced a large quantity of viable 

1974] Genus Kalmia — Ebinger 377 

seed while in others the capsules remained small and lacked 
seeds. From some crosses, both sterile and fertile individu- 
als were produced. Also, among the offspring that pro- 
duced no seeds, stainable pollen varied from 64% to 2%. 

All artificially produced hybrids between these two taxa 
are similar to Kalmia angustifolia var. angustifolia, and 
except for the lack of seed set in some, could not be dis- 
tinguished from it. All individuals have glandular calyces 
and most have leaves that are not puberulent beneath, both 
characteristics of typical var. angustifolia. Only 2 hybrids 
were found that had leaves which were puberulent beneath. 
More studies are needed to clarify the status of these taxa, 
but until more information is available, it seems best to 
maintain them as varieties. 

Kalmia angustifolia L. var. angustifolia f. candida Fern. 
Rhodora 15: 151. 1913. 

Kalmia angustifolia var. candida (Fern.) Rehd. In: 
Bailey, Stand. Cycl. Hort. 3: 1734. 1915. TYPE: Newfound- 
land: damp mossy or turfy hollows on hill south of St. 
John’s, Fernald & Wiegand 6019 (GH). 

According to Jaynes (1971a, 1971b) the presence of pig- 
ment in Kalmia angustifolia is determined by a single 
dominant gene, and the true-breeding recessive is white 
flowered. These white flowered individuals are anthocyan- 
inless with green stems, as contrasted to the normal wild 
types with reddish stems. This characteristic is particularly 
obvious in seedlings. 

It is possible that the recessive genes for the anthocyan- 
inless condition are farily common in populations of this 
species, but are rarely expressed. In a population of Kalmia 
angustifolia from Madison, Connecticut, Jaynes (1971b) 
found that about 24% of the population was heterozygous 
for the anthocyaninless condition, but no homozygous re- 
cessive plants were found. It appears that the anthocyanin- 
less plants are at a selective disadvantage in the wild. 


378 Rhodora [Vol. 76 

(MAINE). LINCOLN CO.: Hyland 69 (MAINE). Massachusetts: Sher- 

born, Loomis (GH). 
CANADA: Newfoundland: St. Johns, Fernald & Wiegand 6019 

4b. Kalmia angustifolia L. var. caroliniana (Small) Fern. 
Rhodora 39: 476, 498. 1937. 
Kalmia caroliniana Small, Fl. Southeast. U.S. 1336. 1903. 
TYPE: North Carolina: Flat Rock, L. R. Gibbs (NY). 
Kalmia carolina Small, Fl. Southeast. U. S. 886. 1903. 
Kalmia angustifolia var. carolina (Small) Fern. Rhodora 
39: 437. 1937. 

Mature leaves permanently puberulent beneath with a 
dense covering of whitish, simple hairs, scattered stalked 
glandular hairs also present; calyx strongly puberulent, 
lacking stalked glandular hairs; corolla usually strongly 
puberulent on the outside. 

Distribution: Open woods and shrubby bogs in the moun- 
tains and sandy woods, pocosins, savannas and bogs on the 
coastal plains. Extending from extreme southern Virginia 
through North Carolina to eastern South Carolina and west 
to eastern Tennessee and northeastern Georgia. 

Common name: Sheep Laurel, Lambkill. 

Georgia: RABUN C0.: Oakey Mt., Duncan 3281 (GH, ID, ISC, MO, 
NY, TENN, US), 4312 (Isc); Tom's Swamp, Bookout & McDowell 65 
(GH). North Carolina: ALLEGHANY CO.: Edmonds, Radford 38191 
(NCU). AVERY CO.: Pineola Bog, Cain & Duncan 251-3 (IND, ND). 
BEAUFORT CO.: Chocowinity, Radford 33396 (NCU). BLADEN CO.: 
White Lake, Kral 27210 (VDB). BUNCOMBE CO.: Asheville, McGarthy 
(US). CARTERET CO.: Beaufort, Bentley (VDB). CHOWAN CO.: Tyner, 
Ahles 39907 (NCU). COLUMBUS C0.: Chadbourn, Wilbur 5315 (DUKE). 
CRAVEN C0.: New Bern, Harbison 14080 (NcU), 14932 (NCU). 
CUMBERLAND CO.: Stedman, Wilbur 5169 (DUKE). DARE CO.: Manns 
Harbor, Radford 8197 (NCU). DUPLIN Co.: Charity, Fogg, Jr. 5481 
(GH, PENN). GATES CO.: Gatesville, Godfrey 5214 (DUKE, GH). 
GREENE CO.: Walstonburg, Radford 36508 (NCU). HARNETT CO.: 
Blomquist 4486 (DUKE). HENDERSON CO.: Muddy Creek, Smith 221 
(180, US). HOKE CO.: Timberland, Ahles 29514 (IND). HYDE CO.: 
Radford & Stewart 803 (NCU). JACKSON CO.: Cullowhee, Biltmore 

1974] Genus Kalmia — Ebinger 379 

Herb. 1344e (ALTA, GH, US). JOHNSTON CO.: Benson, Smith (IND, 
NCU). JONES CO.: Hargetts Store, Radford 36956 (NCU). LENOIR 
co.: Pine Hill, Radford 31651 (NCU). MACON co.: Highlands, 
Wilson 1854 (TENN). MARTIN CO.: Robersonville, Radford 35364 
(Isc). MONTGOMERY CO.: Pekin, Radford 14595 (NCU). NEW HAN- 
OVER CO.: Wilmington, Newber 165 (UARK). ONSLOW co.: Dixon, 
Moldenke 1242 (DUKE, MIN, MO, NY, PENN, US). PASQUOTANK CO.: 
Elizabeth City, Ashe (NCU). PENDER CO.: Wards Corner, Ahles & 
Bell 7856 (MISSA, MSC). PERQUIMANS CO.: Beach Spring, Radford 
5470 (NCU). PITT CO.: Staton, Radford 34940 (NCU). RICHMOND 
co.: Rockingham, Coker (NCU). ROBESON CO.: Allenton, Ahles & 
Ramseur 23770 (ND, PAC). SAMPSON CO.: Tomahawk, Fox & Boyce 
3792 (KY). SCOTLAND CO.: Wagram, Correll 1181 (DUKE). TRAN- 
SYLVANIA CO.: Cedar Mt., Freeman 58130 (NCU). TYRRELL CO.: 
Cross Landing, Radford 33820 (NCU). WASHINGTON CO.: Hoke, 
Radford 35264 (NCU). WATAUGA CO.: Deep Gap, Radford 6239 (NCU). 
WAYNE CO.: Goldsboro, Radford 22006 (NCU). South Carolina: 
DARLINGTON CO.: between Darlington & Society Hill, Smith 1068 
(NCU). HORRY CO.: Radford & Stewart 979 (NCU, NY). ORANGEBURG 
co.: Edisto Swamp, Ravenel (GH). SUMTER CO.: Poinsette St. Park. 
Holdaway (DUKE). Tennessee: JOHNSON CO.: Cole’s Bog, Shanks 
& Sharp 1115 (PENN, TENN); Shady Valley, Barclay (TENN), Sharp 
& Underwood 1252 (vps), 1552 (NCU, NY), 3161 (NY). KNOX CO.: 
Kingston Pike, Jennison 717 (TENN). Virginia: CARROL CO.: Wood- 
lawn, Massey 5058 (GH). NANSEMOND CO.: Whaleyville, Fernald & 
Long 7564 (GH, NY, US). NORFOLK CO.: Dismal Swamp, Coville & 
Kearney, Jr. 91 (08, US), Portsmouth Ditch, Grimes 4517 (MIN, NY). 
SOUTHAMPTON CO.: Franklin, Fernald & Long 10002 (GH). 

The name first applied to this taxon was Kalmia carolina 
when Small (1903) separated the plants with puberulent 
leaves from typical K. angustifolia. Later in a list of genera 
and species in the same publication (page 1336) the name 
was changed to K. caroliniana. It is very possible that this 
later name is a printing error, particularly since the name 
K. carolina was used by the author in a later publication 
(Small, 1914). Fernald (1937) transferred the epithet 
carolina to a variety of K. angustifolia (page 437), but later 
in the same publication (page 476, 498) changed the name 
to variety caroliniana, stating that this spelling, “since it 
accompanied the citation of the type, should prevail.” 

Kalmia angustifolia var. caroliniana has a fairly re- 
stricted range. Most of the specimens examined are from 

380 Rhodora [Vol. 76 

the coastal plain and the mountains of North Carolina with 
the species being extremely common on the pocosins and in 
bogs. This taxon also has a sporadic occurrence in the 
coastal plain of South Carolina (Radford, Ahles, and Bell, 
1968) and in southern Virginia (Fernald, 1938, 1939), 
while it has been reported from a few localities in the moun- 
tains of extreme eastern Tennessee by Jennison (1935) and 
Shanks (1952) and from two mountain bogs in the Blue 
Ridge of northeastern Georgia by Duncan (1948). 

The characteristics used to separate Kalmia angustifolia 
var. caroiimiana are very reliable, but occasionally some 
overlap does occur. Usually, var. angustifolia has glandular 
calyces and leaves that are not puberulent beneath while in 
var. caroliniana the calyces are eglandular and the leaves 
are strongly puberulent beneath. All but two specimens of 
var. caroliniana examined had strongly puberulent leaves 
and even those were lightly puberulent on the undersurface 
of the leaf. This was also noticed by Radford, Ahles, and 
Bell (1968) who mentioned that a few specimens of var. 
caroliniana with glabrate leaves were found in the Caro- 
linas. A few specimens of var. angustifolia from Nantucket 
Island and the coast of Massachusetts and Connecticut have 
leaves that are strongly puberulent beneath. All of these, 
however, have strongly glandular calyces, typical of var. 
angustifolia, and are referred to that taxon. The best char- 
acter to separate the two varieties is the presence or absence 
of stalked glandular hairs on the calyx. Close examination, 
however, reveals that on some specimens of var. caroliniana 
a few glandular hairs are present on the calyces. The glands 
are not very numerous, however, and mostly restricted to 
the margins of the lobes or to the base of the calyx. Even 
on the holotype a few of these stalked glandular hairs are 
found. Because of this variability it is probably best to 
maintain these taxa as varieties of one species rather than 
as separate species. 

A white flowered, anthocyaninless form of var. carolin- 
iana also exists. It is similar genetically to Kalmia angusti- 
folia var. angustifolia f. candida in that it is a homozygous 

1974] Genus Kalmia — Ebinger 381 

recessive plant in which flower color is determined by a 
single pair of genes (Jaynes, 1971b). A specimen of this 
form was obtained by Dr. Jaynes from the Garden-in-the- 
Woods, in Sudbury, Massachusetts. No wild individuals 
have been found and its origin is not known. The best 
crossing results obtained by Jaynes (1968a) between K. 
angustifolia and other species of the genus were when this 
white flowered form was used as the female parent (Jaynes, 

5. Kalmia cuneata Michaux, Fl. Bor.-Amer. 1: 257. 1803. 
Chamaedaphne cuneata (Michx.) Kuntze, Rev. Gen. Pl. 
2: 388. 1891. TYPE: In Carolina, Michaux s.n., (GY). 

Much branched rhizomatous shrub to 1.5 m tall; branch- 
lets terete, reddish brown, usually adscending, stipitate- 
glandular and puberulent, becoming glabrous and light 
gray; leaves alternate, deciduous, flat, oblanceolate to ellip- 
tic-lanceolate, 2-6 cm long, 0.5-1.5 cm wide, lightly stipitate 
glandular beneath, glabrous above, apex obtuse to acute, 
apiculate, base narrowly cuneate, midrib puberulent above, 
stipitate-glandular beneath; petiole to 4 mm long; inflor- 
escence a reduced corymbiform raceme or fascicle of 3-10 
flowers in the upper axils of the last season’s growth, pedi- 
cels 1-3 mm long, glabrous to lightly stipitate-glandular, 
recurved in fruit, subtended by 2 bracteoles in the axil of a 
small lanceolate bract about 1 mm long, main axis of the 
inflorescence less than 3 mm long; calyx glabrous to lightly 
stipitate-glandular, 6-7.5 mm broad, the lobes oblong, apex 
acute; corolla creamy white with a red band within, lightly 
stipitate-glandular outside, puberulent near the base within, 
1.3-1.8 mm broad; anthers purplish brown, 1-1.3 mm long; 
filaments puberulent toward the base; style 7-10 mm long; 
ovary stipitate-glandular; capsule depressed globose, 4-6 
mm broad, 2.5-3.5 mm long, stipitate-glandular; seeds light 
brown, 0.6-0.9 mm long, the testa not loose or extended past 
the ends of the seed. Chromosome number n — 12. 

Distribution: Wet thickets and shrub bogs in southeastern 

382 Rhodora [Vol. 76 

North Carolina and adjacent South Carolina. 
Common name: White Wicky. 


North Carolina: BLADEN CO.: Colly Creek, Ashe (PH); 10 miles 
north of Elizabethtown, Ebinger 11271 (EIU); Jerome Pocosin, Buell 
& West 1740 (MIN). CUMBERLAND C0.: Hope Mill, Alexander (NY). 
MOORE CO.: Pinehurst, Wicker (NY). PENDER CO.: Squalling Bluff 
Rd., Coville (MICH, NY, US); Burgaw, Chrysler 2262 (CHRB). RICH- 
MOND CO.: Hamlet, Wiegand & Manning 2365 (GH). South Carolina: 
DARLINGTON CO.: Hartsville, Smith (NY), Coker (NY). 

A small fragment of the type of Kalmia cuneata is at the 
Gray Herbarium. It consists of one fascicle of 5 flowers and 
a small stem with a few leaves attached. On the label is 
written “Ex Herb. Michaux Kalmia cuneata." At the pres- 
ent time the rest of the type has not been located. 

Kalmia cuneata is very distinct. The only species that it 
could possibly be confused with is K. angustifolia since 
their general habit and leaf size are similar. Kalmia cune- 
ata, however, is the only deciduous member of the genus 
and the only one in which the flowers are borne in fascicles 
from the axils of last season’s growth. Other distinguishing 
characteristics are the creamy-white flowers with a red 
band within, the oblanceolate leaves, and the recurved fruit- 
ing pedicels. 

Genetically, this species appears to be most closely re- 
lated to Kalmia hirsuta, another alternate leaved species of 
the southeastern United States. Jaynes (1968a) reported 
that K. cuneata set no seed when used as the female parent, 
but as the male parent crossed with both K. angustifolia 
and K. hirsuta. The cross with K. angustifolia produced 
only inviable hybrids. Most of the hybrids with K. hirsuta 
were also inviable, producing sectors of albino or yellow- 
green tissue, but one hybrid flowered and was pollen sterile. 

This species is one of the rarest shrubs in North America 
and few early reports exist concerning its distribution. 
Pursh (1814) reports that it occurs on the mountains of 
Carolina, while Nuttall collected it between Camden, South 
Carolina and Statesville, North Carolina (Wood, 1961). 

1974] Genus Kalmia — Ebinger 383 

Curtis (1835) reported its occurrence about 30 miles NE 
of Wilmington, North Carolina. Present studies indicate 
that it grows only in the sandhills and bay margins of the 
pocosin ecotones of the Carolinas. It now is known from 
8 counties in this region (Radford, Ahles & Bell, 1968). 

6. Kalmia hirsuta Walt. Fl. Carol. 138. 1788. 
Chamaedaphne hirsuta (Walt.) Kuntze, Rev. Gen. Pl. 2: 
388. 1891. Kalmiella hirsuta (Walt.) Small, Fl. Southeast. 
U.S. 886. 1903. TYPE: unknown. 
Kalmia ciliata Bartram, Trav. N. & S. Carol. v, 7, 94, 171, 
242,1791. (nom. nud.). 

Low, sparingly branched shrub, to 6 dm tall with a thick 
basal caudex; branchlets terete, reddish brown, puberulent 
and densely hispid; leaves alternate, sessile or with a petiole 
to 1 mm long, elliptic to ovate, 5-14 mm long, 1.5-8 mm 
broad, the margins usually slightly revolute, usually pu- 
berulent and hispid on both surfaces, small stipitate glands 
usually present beneath, apex acute, base broadly cuneate 
to rounded; flowers solitary (rarely in fascicles of 2-5 or 
in compact racemes) in the axils of the leaves of new 
growth; pedicels to 1.5 cm long, stipitate-glandular and 
hispid, subtended by a pair of bracts; calyx green, the lobes 
foliaceous, lanceolate, 3-8 mm long, hispid, tardily deciduous 
in fruit; corolla pink (rarely white), marked with red 
around the anther pockets and a red ring near the base, 
1-1.5 em broad, usually lightly hirsute and stipitate-glandu- 
lar on the keels outside, puberulent at the base within; 
anthers purplish brown, 0.7-1.3 mm long; filaments puberu- 
lent toward the base; ovary usually stipitate glandular; 
style 5-7 mm long; fruit 2-4 mm broad, 2-3.5 mm long, 
subglobose to ovoid, thinly stipitate-glandular; seeds light 
brown, 0.3-0.5 mm long, the testa not loose or extended past 
the ends of the seed. Chromosome number n = 12. 

Distribution: Low, sandy pine savanna, sand hills, dunes 
and flat pine woods on the coastal plains of southern Ala- 

384 Rhodora [Vol. 76 

bama and Georgia, northern Florida and the southern tip 
of South Carolina. 
Common name: Sandhill Laurel, Calico-bush. 

Alabama: BALDWIN CO.: Gulf State Park, Webster & Wilbur 3539 
(GH, NY, US). MOBILE C0.: Theodore, Pennell 4513 (MICH, NY, PENN, 
PH). Florida: ALACHUA CO.: Cross Creek, Ford 2250 (GH, TENN). 
BAY CO.: Lynn Haven, Billington (MicH, us); Lazor 5006 (Pac). 
CLAY CO.: Kingsley Lake, Henry 3828 (PH). COLUMBIA CO.: Lake 
City, Nash 2193 (GH, MASS, MICH, MIN, MSC, ND, NY, OS, PH, US). 
DIXIE C0.: Old Town, West & Arnold (PH). DUVAL CO.: Jacksonville, 
Curtis 4359 (ARIZ, ND, NY, US), 4921 (GH, ISC, MIN, MSC, NY, US). 
ESCAMBIA CO.: Pensacola, Webster & Wilbur 3572 (GH, NY, US). 
FLAGLER CO.: Bunnell, Ames (GH). FRANKLIN CO.: Carrabelle, Mosier 
884 (MIN, NY, PENN, US); Moldenke 1134 (MIN, NY, PENN, US). 
GADSDEN CO.: Quincy, Sargent (MIN). GILCHRIST CO.: Trenton, Ward, 
West & Will 2651 (GH, US). GULF CO.: Port St. Joe, Ward & Ward 
1460 (GH). HAMILTON CO.: Bowers & Wofford 71-550 (TENN). 
HOLMES CO.: Bonifay, Blanton 6582 (US). JACKSON CO.: Compass 
Lake, Knight & Arnold (PAC). JEFFERSON Co.: Wacissa, Godfrey 
53552 (GH). LAFAYETTE CO.: Smith & Myint 189 (GH). LEON CO.: 
Silver Lake, Godfrey 53660 (GH, MISSA, NY). LEVY CO.: Bronson, 
Godfrey & Lindsey 56989 (GH). LIBERTY CO.: Bristol, Small, Small 
& DeWinkeler 11448 (Gn, NY, PENN, TENN). MADISON CO.: Green- 
ville, Ramsey 106 (MSC). MARION CO.: Ocala, Beardslee, Jr. (MU). 
OKALOOSA CO.: Crestview, McArthur (GH). PUTNAM CO.: Interlachen, 
Fox 5703 (GH, MICH, MIN, NY, TENN). ST. JOHNS CO.: Anastasia 
Island, Crawford (NY, PH). ST. LUCIE CO.: Fort Walton, Henry 
1087 (PH). SANTA ROSA CO.: Munson, Ford & Arnold 4190 (us). 
WAKULLA CO.: St. Marks Wildlife Refuge, Godfrey 57860 (GH). 
WALTON CO.: Santa Rosa, Ripley & Barneby 10128 (NY). Georgia: 
APPLING CO.: Baxley, Cottam 11060 (UT); Ebinger 11204 (E10). 
BRANTLEY CO.: Waynesville, Ebinger 11232 (EIU). BULLOCH CO.: 
Portal, Henry 7311 (PH). CAMDEN CO.: Waverly, Ebinger 11228 
(EIU). CANDLER CO.: Metter, Henry 7277 (PH); Hermann 10122 
(GH, US). CHARLTON CO.: Folkston, Harbison 5816 (GH). COFFEE 
co.: Harper 687 (GH, US). EFFINGHAM CO.: Harper 961 (GH). 
GLYNN C0.: Emanuel Church Rd., Ebinger 11231 (EIU). JEFF DAVIS 
cc.: Hazlehurst, Ebinger 11195 (EIU). LANIER CO0.: Adams 50 (Ky). 
LIBERTY CO.: Altamaha River, Small (GH, MIN, NY, PH). LOWNDES 
co.: Valdosta, Beardslee, Jr. (MU). MCINTOSH CO.: Darien, Cron- 
quist 5366 (GH, MICH, NY, PH). TATTNALL CO.: Reidsville. Cronquist 
5840 (GH, IND, MICH, NY, PH, US). THOMAS CO.: Thomasville, Taylor 
(GH, MIN): TIFT CO.: Tifton, Hardin & Duncan 13649 (US). WARE 

1974] Genus Kalmia — Ebinger 385 

co.: Suwannee Lake, Harper 48 (PH), 298 (GH, NY, PH, US). WAYNE 
co:. Grangerville, Ebinger 11233 (EIU). WORTH CO.: Poulan, Pollard 
& Maxon 563 (NY, Us). South Carolina: BEAUFORT CO.: Ferry 
Landing, Bell 3805 (NY). HAMPTON CO.: Hampton, Leonard & Rad- 
ford 1662 (ARIZ, BRY, CHRB, NHA). JASPER CO.: Ridgeland, Ebinger 
11242 (EIU). 

Except for transfers of the specific epithet, hirsuta, to 
other genera, only one other name has been proposed for 
this taxon. Bartram (1791) used Kalmia ciliata for a plant 
he observed on his travels through the southeastern United 
States. He used this name a number of times but did not 
include a description (Merrill, 1945). He did, however, 
describe “a diminutive but elegant Kalmia" (page 18) 
which undoubtedly refers to K. ciliata. This description 
leaves no doubt that his K. ciliata is the same as K. hirsuta. 
The first time the name K. ciliata appears with a descrip- 
tion is in the work of Dumont de Courset (1811) who 
attributes the name to Bartram. 

Kalmia hirsuta has a relatively limited distribution, oc- 
curring along the coastal plain in southeastern United 
States. Small (1903, 1914) indicated that it extended from 
Virginia to Florida, perpetuating an error started by Gray 
(1856). Its occurrence in Virginia is extremely doubtful 
(Fernald, 1947) and is probably due to a few herbarium 
specimens with incomplete or erroneous collecting data. 
Two such specimens have been found during the present 
study. Both were collected during the middle of the last 
century and beside the collector’s name and the year col- 
lected include the name Virginia. 

Most of the specimens of Kalmia hirsuta examined are 
hispid on the stems, leaves, and pedicels. A few specimens 
were found, however, that were completely glabrous, lack- 
ing the long coarse hairs as well as the stipitate-glands and 
smaller hairs commonly found on most parts of the plant. 
Variation also occurs in the arrangement of the flowers on 
the plant. Usually the flowers are solitary in the axils of 
the leaves, but occasionally the flowers occur in fascicles of 
2-5, while a few specimens were found in which short 

386 Rhodora [Vol. 76 

racemes occur in the axils of the leaves. In other specimens 
the flowers appeared as a terminal raceme due to a shorten- 
ing of the internodes and a reduction in the size of the sub- 
tending leaves. The pedicels also vary extensively in length. 
The flowers of some plants are nearly sessile, while in others 
the pedicels are 1.5 cm long with the two basal bracts 
scattered along the stalk. 

Genetically Kalmia hirsuta appears to be closely related 
to K. latifolia with which it can be crossed to produce viable 
hybrids with 0-10% pollen stainability. Jaynes (1968a) also 
reported that it can be crossed with both K. angustifolia 
and K. cuneata producing viable but sterile hybrids. Based 
on habit and morphology K. hirsuta is most closely related 
to K. ericoides. These species were placed in the genus 
Kalmiella by Small (1914) since they have a deciduous 
calyx and their flowers are borne singly in the axils of the 
leaves. Unfortunately the Cuban member of the genus is 
not available for genetic studies. 

7. Kalmia ericoides Wright ex Griseb. Cat. Pl. Cubensium 
51. 1866. 

Chamaedaphne ericoides (Wright ex Griseb.) Kuntze, 
Rev. Gen. Pl. 2: 388. 1891. Kalmiella ericoides (Wright ex 
Griseb.) Small, North Amer. Fl. 29: 54. 1914. TYPE: Cuba 
occ. in distr. Guane, in pinetis arenosis pr. La Grifa, C. 
Wright 2199 (GH, NY, US - Isotypes). 

A sparingly branched, erect to spreading shrub, to 10 dm 
tall with a thick basal caudex; branchlets light gray to 
reddish-gray, puberulent and commonly densely hispid and 
stipitate-glandular (rarely glabrous) ; leaves alternate, per- 
sistant, thick, leathery, subsessile or with a puberulent 
petiole to 1 mm long, linear, 4-13 mm long, 0.6-2 mm broad, 
the margins strongly revolute, usually lightly hispid and 
stipitate-glandular on both surfaces, sometimes strongly 
puberulent (rarely glabrous); flowers solitary (rarely in 
fascicles or in compact racemes) in the axils of the leaves 
at the end of the branches, forming a terminal cluster; 
pedicels to 13 mm long, commonly hispid and stipitate- 

1974] Genus Kalmia — Ebinger 387 

glandular, subtended by a pair of puberulent bracts 1-3 mm 
long; calyx green, the lobes foliaceous, lanceolate, 3-6 mm 
long, usually hispid and stipitate-glandular, tardily decidu- 
ous in fruit; corolla light pink to white, 10-17 mm broad, 
usually lightly hirsute and stipitate-glandular on the keels 
outside, lightly puberulent at the base within; anthers pur- 
plish brown, 0.8-1.2 mm long; filaments puberulent toward 
the base; ovary usually stipitate-glandular; style 4-7 mm 
long; fruit 2-4 mm broad, 2-3 mm long, subglobose to ovoid, 
lightly stipitate-glandular ; seeds reddish-brown, 0.4-0.7 mm 
long, the testa not loose or extending past the ends of the 

Kalmia ericoides is the only member of the genus that 
has not been studied cytologically. Morphologically, how- 
ever, it is most closely related to K. hirsuta, a species re- 
stricted to southeastern United States. Both have relatively 
small leaves (less than 1.5 em long), that are usually hispid, 
their flowers are borne singly in the axils of the leaves, and 
the calyx is tardily deciduous. The two are easily separated, 
however, since the leaves of K. ericoides are thick and 
leathery with revolute margins while in K. hirsuta the 
leaves are thin and lack a strong revolute margin. Also, 
the flowers are borne scattered along the stem in the axils 
of the leaves in K. hirsuta while in K. ericoides, they are 
solitary in the leaf axils, but clustered toward the ends of 
the stems giving the appearance of a terminal corymbiform 

Kalmia ericoides is endemic to savannas and pine barrens 
of western Cuba. Although it has a very limited distribu- 
tion, the variation that exists in pubescence and compact- 
ness of the inflorescence has resulted in a division of this 
complex into three species by Alain (1957), while others 
considered it a single highly variable species (Wood, 1961). 
Present studies indicate that the compactness of the inflor- 
escence is not completely reliable in separating the taxa. 
The variation in leaf pubescence, however, is relatively 
stable. As a result, the following two varieties are recog- 

888 Rhodora [Vol. 76 


1. Leaves glabrous or lightly hispid and with a few scat- 
tered stalked glandular hairs on the upper surface. .... 
rr rr Ta. var. ericoides. 

1. Leaves densely puberulent and stipitate-glandular on 
the upper surface. .............. 7b. var. aggregata. 

Ta. Kalmia ericoides Wright ex. Griseb. var. ericoides. 

Kalmiella simulata Britt. & Wils. Mem. Torrey Bot. Club 
16: 93. 1920. TYPE: Cuba: vicinity of Los Indios, Isle of 
Pines, Britton, Britton, & Wilson 14205 (NY). 

Shrub with ascending, hispid, stipitate-glandular and 
sometimes puberulent branchlets; leaves scattered, usually 
only 5-12 per cm of branchlet, glabrous to lightly hispid and 
stipitate-glandular above, not puberulent, 4-9 mm long, 
0.6-1.5 mm wide; pedicels usually longer than the subtend- 
ing leaves, giving the inflorescence an open appearance; 
calyx lobes 3-4.5 mm long, hispid and lightly puberulent on 
the outside, usually glabrous within. 

Distribution: Low, white-sand savanna and pine barrens 
on the Isle of Pines and in the Province of Pinar del Rio, 


Cuba: PINAR DEL RIO: La Grifa, Ekman 18165 (NY, Us), Wright 
2199 (GH, NY, US); Laguna de Alcatraz Grande, León & Victorin 
18706 (GH, US), 20474 (GH, Us); El Sabalo, Alain 1326 (GH, US). 
ISLE OF PINES: Los Indios, Britton Britton, & Wilson 14205 (GH, 
NY), Killip 42882 (GH, NY, US), León & Victorin 17853 (GH), Vic- 
torin & Alain TTa (US). 

Kalmia ericoides var. ericoides has a very restricted range 
and few herbarium specimens are available. It appears to 
be a fairly stable variety that varies mostly in the extent of 
leaf pubescence. All of the specimens seen from the prov- 
ince of Pinar del Rio are typical of the type specimen which 
has a few scattered glandular hairs and is lightly hispid on 
the upper leaf surface. In contrast, most of the specimens 
of this variety from the Isle of Pines are glabrous on the 

1974] Genus Kalmia — Ebinger 389 

upper leaf surface. These glabrous specimens were sepa- 
rated as Kalmiella simulata by Britton and Wilson (Britton, 
1920). These glabrous specimens all have small, narrow 
leaves (less than 9 mm long and 0.6-1.2 mm wide), pedicels 
that are much longer than the subtending leaves, and calyx 
lobes that are fairly short (less than 4.5 mm long) all char- 
acteristics of variety ericoides. 

7b. Kalmia ericoides var. aggregata (Small) Ebinger, 
comb. nov. 
Kalmiella aggregata Small, North Amer. FI. 29: 54. 1914. 
Kalmia aggregata (Small) Copeland, Amer. Midl. Nat. 30: 
571. 1943. TYPE: Cuba: pinelands, Los Indios, Isle of Pines, 

Jennings 324 (NY). 

Shrub with adscending, lightly hispid and strongly pu- 
berulent branchlets; leaves crowded, usually 15-25 per cm 
of branchlet, densely puberulent and stipitate-glandular 
above, 8-13 mm long, 1-2 mm wide; pedicels usually as short 
as the subtending leaves, giving the inflorescence a compact 
appearance; calyx lobes 3.5-5.5 mm long, hispid and densely 
puberulent on the outside, densely puberulent within. 

Distribution: Low, white-sand savanna and pine barrens 
on the Isle of Pines, Cuba. 


Cuba: ISLE OF PINES: Beach at Siquanea, Morton 10098 (us); 
between Mina de Oro and Playa del Soldado, Killip 43905 (us), 
45385 (GH); Lama Daquille, Ekman 72492 (NY); Los Indios, Jen- 
nings 324 (GH, NY, US), León & Victorin 18857 (GH, US), Victorin 
& Alain 77 (GH, NY, US), Playa Roja, Killip 48001 (GH, NY, US); 
San Pedro, Britton, Britton, & Wilson 14146 (GH, NY, US), Killip 
43698 (US), 45566 (vs), Morton 10018 (Us), 10053 (US); Siquanea 
region, Killip 44043 (us), 44044 (us), 44753 (us), 44755 (US). 

This variety appears to be more robust than the preced- 
ing. Specimens to 13 dm have been reported, and the leaves 
and flowers are, in general, slightly larger than in variety 
ericoides. Besides this difference in size, all of the speci- 
mens of variety aggregata examined are strongly puberu- 
lent throughout. In particular the leaves are strongly 

390 Rhodora [Vol. 76 

puberulent on both surfaces, but this is also characteristic 
of the branches, pedicels and sepals. 


Kalmia lanceolata Raf. Autikon Bot. 86. 1840. This 
name should probably be referred to Kalmia polifolia but 
the description could apply to other taxa. No specimen to 
represent this name has been found. 

Kalmia serotina Hoffmgg. Verz. Pflanzenkulturen 70. 
1824. Nomen nudum. 

Kalmia spuria Bartr. Trav. N. & S. Carol. 308. 1791. 
Nomen nudum. 


ABRAMS, L. 1951. Illustrated Flora of the Pacific States: Wash- 
ington, Oregon, and California, Vol. 3. Stanford. 

AITON, W. 1789. Hortus Kewensis. Vol. 2. London. 

ALAIN, BR. H. 1957. Ericaceae. In: LEON, BR. H. & BR. H. ALAIN, 
Flora de Cuba. Vol. 4. Havana. 

AMERICAN FORESTRY ASSOCIATION. 1955. These are the Champs. 
Am. Forests 61(9): 31-40. 

ANONYMUs. 1895. New or little-known plants. Garden Forest 8: 

ASHLEE, T. R. 1932. A contribution to the Latah flora of Idaho. 

Northw. Sci. 6: 69-82. 

BARTON, B. S. 1802. Some account of the poisonous and injurious 

honey of North America. Trans. Am. Phil. Soc. 5: 51-70. 
1804. Collections for an essay toward a materia medica 

of the United States. Philadelphia. 

BaRTRAM, W. 1791. Travels through North and South Carolina, 
Georgia, East and West Florida. Philadelphia. 

BEAL, W. J. 1867. Agency of insects in fertilizing plants. Am. 
Nat. 1: 254-260. 

BEAN, W. J. 1897. Kalmias. Garden 52: 77-78. 

BEISSNER, L., E. SCHELLE, & H. ZABEL. 1903. Handbuch der 
Laubholz-Benennung. Berlin. 

Benson, A. B. 1937. Peter Kalm’s travels in North America. New 

Berry, E. W. 1907. Contributions to the Mesozoic flora of the 
Atlantie coastal plain. II. North Carolina. Bull. Torrey Bot. 
Club 34: 185-206. 

1974] Genus Kalmia — Ebinger 391 

1913. Contributions to the Mesozoic flora of the At- 
lantie coastal plain. IX. Alabama. Bull. Torrey Bot. Club 
40: 567-574. 

1919. Upper Cretaceous floras of the eastern gulf 
region in Tennessee, Mississippi, Alabama, and Georgia. U. S. 
Geol. Surv. Prof. Pap. 112: 1-177. 

1920. Contributions to the Mesozoic flora of the At- 
lantic coastal plain. XII. North Carolina. Bull. Torrey Bot. 
Club 47: 397-406. | 

, & W. A. JouNSTON. 1922. Pleistocene interglacial 
deposits in the Vancouver region, British Columbia. Proc. Trans. 
Roy. Soc, Canada 16(4): 133-140. 

BrGELOW, J. 1818. American medical botany.  1(II): 133-141; 
3(II): 185-186. Boston. 

BorviN, B. 1967. Connecting vowels in epithets of Latin origin. 
Rhodora 69: 451-455. 

1968. Flora of the prairie provinces. A handbook to 
the flora of the provinces of Manitoba, Saskatchewan and 
Alberta. Phytologia 16: 1-47. 

BRITTON, E. G. 1913. Wild plants needing protection. Jour. N. Y. 
Bot. Gard. 14: 121-123. 

1925. A freak of the mountain laurel. Ibid. 26: 187- 

Britton, N. L. 1908. North American trees. New York. 

1920. Description of Cuban plants new to science. 
Mem. Torrey Bot. Club 26: 57-118. 

BRowN, R. W. 1962. Paleocene flora of the Rocky Mountains and 
Great Plains. U. S. Geol. Surv. Prof. Pap. 375: iv + 119 + 69. 

BROWNE, D. J. 1857. The trees of America. New York. 

BRUNET, L. O. 1867. Catalogue des végétaux ligneux du Canada. 
BUTTRICK, P. L. 1924. Connecticut's state flower, the mountain 
laurel, a forest plant. Marsh Bot. Gard. Publ. 1: 1-28. 
CALLAN, H. G. 1941. The cytology of Gaulthettya wisleyensis 
(Marchant) Rehder. A new mode of species formation. Ann. 
Bot. II. 5: 579-585. 

Cary, C. A, & F. G. MATTHEWS. 1903. Some diseases of cattle. 
Alabama Agr. Exp. Sta. Bull. 125: 105-168. 

CaTEsBY, M. 1743. The natural history of Carolina, Florida and 
the Bahama Islands. London. 

CHENON, L. J. 1751. Dissertation Botanica, qua Nova Plantarum 
Genera. Upsaliae. 

CLAWSON, A. B. 1933. Alpine Kalmia (Kalmia microphylla) as a 
stock-poisoning plant. U. S. Dept. Agr. Tech. Bull. 391: 1-9. 

392 Rhodora [Vol. 76 

COLVIN, W. S., & W. S. EISENMENGER. 1943. Relationships of natu- 
ral vegetation to the water holding capacity of the soils in New 
England. Soil Sci. 55: 433-446. 

CoPELAND, H. F. 1943. A study, anatomical and taxonomic, of the 
genera of Rhododendroideae. Am. Midl. Nat. 30: 533-625. 

Cox, H. T. 1948. Studies in the comparative anatomy of the 
Ericales. I.  Ericaceae — Subfamily Rhododendroideae. Am. 
Midl. Nat. 39: 220-245. 

CRAWFORD, A. C. 1908. Mountain laurel, a poisonous plant. U. S. 
Dept. Agr. Bur. Pl. Ind. Bull. 121: 21-35. 

Curtis, M. A. 1835. Enumeraticn of plants growing spontaneously 
around Wilmington, North Carolina, with remarks on some new 
and obscure species. Boston Jour. Nat. Hist. 1: 82-141. 

CurTis, W. 1795. Kalmia angustifolia. Narrow-leaved Kalmia. Bot. 
Mag. 9: 331. 

Davis, P. H., & V. H. HEvwoop. 1963. Principles of Angiosperm 
taxonomy. Princeton. 

DEAM, C. C. 1940. Flora of Indiana. Indianapolis. 

DEANE, W. 1899. Kalmia latifolia in Vermont. Rhodora 1: 136. 

DEGRAFFE, B. L. 1896. The tannins of some Ericaceae. Am. Jour. 
Pharm. 68: 313-321. 

DRUDE, O. 1889. Ericaceae. In: ENGLER, A., & K. PRANTL, Die 
naturlichen Pflanzenfamilien. IV(1): 15-65. 

DuprEY, T. R. 1967. Ornamental mountain-laurel and a new cul- 
tivar: Kalmia latifolia ‘Bettina’. Am. Hort. Mag. 46: 245-248. 

DUMONT DE CoursET, G. L. M. 1802. Le botaniste cultivateur. 
Vol. 2. Paris; 1811. Ed. 2, Vol. 3. Paris. 

Duncan. W. H. 1948. Preliminary reports on the flora of Georgia. 
I. The distribution in Georgia of Spermatophytes new to or 
rare in the state. Castanea 13: 70-83. 

EGLER, F. E. 1947. Effects of 2,4-D on woody plants in Connecti- 
cut. Jour. Forest. 45: 449-452. 

—. 1948. 2.4-D effects in Connecticut vegetation, 1947. 

Ecology 29: 382-386. 
1949. Herbicide effects in Connecticut vegetation, 1948. 

Ecology 30: 248-256. 
. 1950. Herbicide effects in Connecticut vegetation, 1949. 

Bot. Gaz. 112: 76-85. 

ENGELMANN. G.. W. M. CANBY. & A. GRAY. 1877. Large trunks of 
Kalmia latifol'a. Amer. Nat. 11: 175. 

FERN^'Dp. M. L. 1911. Notes from the Phaenogamic Herbarium of 
the New England Botanical Club. II. Rhodora 13: 177-183. 

1937. Local plants of the inner coastal plain of south- 

eastern Virginia. Rhodora 39: 321-366, 379-415, 433-459, 465-491. 

. 1938. Noteworthy plants of southeastern Virginia. 
Rhodora 40: 434-459. 

1974] Genus Kalmia — Ebinger 393 

1939. Last survivors in the flora of tidewater Vir- 
ginia. Rhodora 41: 529-559. 
1940. A new southern Kalmia. Rhodora 42: 53-54. 

— — — —— 1947. Additions to and subtractions from the flora of 
Virginia. Rhodora 49: 145-159. 

1950. Gray's Manual of Botany. 8th ed. New York. 

FLEMING, C. E. 1920. Project 22 — Poisonous range plants. Nevada 
Agr. Exp. Sta. Ann. Rep. 1920: 39-43. 

FLEMER, W. 1949. The propagation of Kalmia latifolia from seed. 
Bull. Torrey Bot. Club 76: 12-16. 

Forges, E. B. & S. I. BECHDEL. 1930. The chemical composition 
and feeding value of Laurel and Rhododendron, and the condi- 
tions of browsing in forests overstocked with deer, in relation 
to the death of young deer. Penn. Agr. Exp. Sta. Bull. 258: 

& .1931. Mountain Laurel and Rhododendron 
as food for the white tailed deer. Ecology 12: 323-333. 

FULLER, G. D. 1944. Notes on some plants that are rare in Illinois. 
Trans. Ill. St. Acad. Sci. 37: 57-58. 

GLEASON, H. A. 1952. The new Britton and Brown Illustrated 
Flora of the Northeastern United States and adjacent Canada. 
Vol. 3. Lancaster. 

GRAY, A. 1856. Manual of the Botany of the Northeastern United 
States. 2nd ed. New York. 

1871. Dialysis with staminody in Kalmia latifolia. 
Amer. Nat. 1: 373-374. 

1876. How flowers are fertilized. Amer. Agr. 35: 262. 

1879. Gray’s Botanical Text-book. 6th ed. Vol. 1. 
Structural Botany. New York. 

GRIFFITH, R. E. 1847. Medical Botany. Philadelphia. 

Griscom, L. 1931. Dwarf mistletoe and other plants new to New 
Jersey. Rhodora 33: 101. 

Gronovius. J. F. 1739. Flora Virginica. Leiden. 

Gussow. W. C. 1933. Contributions to the knowledge of the flora 
of Northern Manitoba and the North-western Territories, Do- 
minion of Canada (Lat. 58°-62°; Long. 95°-100°. Hudson Bay). 
Canad. Field-Nat. 47: 116-119. 

HAGERUP, O. 1928. Morphological and cytological studies of Bi- 
cornes. Dansk Bot. Ark. 6: 1-26. 

HALL, H. M. 1912. New and noteworthy California plants, I. 
Univ. Calif. Publ. Bot. 4: 195-208. 

HARDIN. J. W., & J. M. ARENA. 1969. Human poisoning from native 
and cultivated plants. Durham. 

HASSKARL, J. C. 1863. Ueber Kalmia latifolia L. Bot. Zeit. 31: 

394 Rhodora [Vol. 76 

Hopepon, A. R. 1955. Ilex glabra and a new station for Kalmia 
latifolia in New Hampshire. Rhodora 57: 34-36. 

HOLLICK, A. 1892. Additions to the Paleobotany of the Cretaceous 
formation of Staten Island. Trans. N. Y. Acad. Sci. 12: 28-39. 

1906. The Cretaceous flora of southern New York and 
New England. U. S. Geol. Surv., Monogr. 50: 1-219. 

1931. Plant remains from a Pleistocene lake deposit 
in the Upper Connecticut River Valley. Brittonia 1: 35-56. 

HorLMEs, E. M. 1884. Medical plants used by the Cree Indians. 
Hudson's Bay territory. Am. Jour. Pharm. 56: 617-621. 

HornMES, M. L. 1956. Kalmia, the American laurels. Baileya 4: 

Howes, F. N. 1949. Sources of poisonous honey. Kew Bull. 1949: 

HULTÉN, E. 1948. Flora of Alaska and Yukon. Vol. 8. Lund. 

1968. Flora of Alaska and neighboring Territories. 

IkUusE, M. 1954. The presence of the viscid threads among pollen 
grains in Phyllodoceae, etc. of Ericaceae. Jour. Jap. Bot. 29: 

JacoBS, M. L, & W. R. Lrovp. 1939. A phytochemical study of 
Kalmia angustifolia. Jour. Am. Pharm. Assoc. 28: 408-412. 
JAYNES, R. A. 1968a. Interspecific crosses in Kalmia. Am. Jour. 

Bot. 55: 1120-1125. 
1968b. Self incompatibility and inbreeding depression 
in three laurel (Kalmia) species. Proc. Amer. Soc. Hort. Sci. 
93: 618-622. 
1968c. Breaking seed dormancy of Kalmia hirsuta 
with high temperatures. Ecology 49: 1196-1198. 
1969. Chromosome counts of Kalmia species and re- 
valuation of K. polifolia var. microphylla. Rhodora 71: 280-284. 
1971a. Laurel selections from seed. True breeding red- 
budded mountain laurel. Cire. Conn. Agr. Exp. Sta. 240: 1-10. 
1971b. A gene controlling pigmentation in sheep laurel. 
Jour. Hered. 62: 201-203. 
1971c. Seed germination of six Kalmia species. Jour. 
Am. Soc. Hort. Sci. 96: 668-672. 
. 1971d. The Kalmias and their hybrids, Quart. Bull. 
Am. Rhododendron Soc. 25: 160-164. 
. 1972. Selection and propagation of improved Kalmia 
latifolia cultivars. Int. Pl. Prop. Soc. 21: 366-373. 

JENNISON, H. M. 1935. Notes on some plants of Tennessee. Rhodora 
37: 309-323. 

JOHNSON, E. A., & J. L. KovNER. 1956. Effects on stream flow of 
cutting a forest understory. Forest Sci. 2: 82-91. 

1974] Genus Kalmia — Ebinger 395 

KENNEDY, G. W. 1875. Arbutin in Kalmia latifolia, Lin. Am. 
Jour. Pharm. 47: 5-6. 

KiNGSBURY, J. M. 1964. Poisonous plants of the United States and 
Canada. Englewood Cliffs. 

KRAUSEL, R., & H. WEYLAND. 1959. Kritische Untersuchungen zur 
Kutikularanalyse Tertiàrer blátter. IV. Dikotyledonen aus der 
braunkohle. Palaeontographica Abt. B. 105: 101-124. 

KUNTZE, O. 1891. Revisio Generum Plantarum. Part II. Leipzig. 

KunMES, E. A. 1961. The ecology of mountain laurel in southern 
New England. Ph.D. Thesis, Yale Univ. 

1967. The distribution of Kalmia latifolia L. Am. 
Midl. Nat. 77: 525-526. 

LAMSON-SCRIBNER, F. 1891. A sketch of the flora of Orono, Me. 
Bot. Gaz. 16: 228-234. 

LANJOUW, J. 1966. International code of botanical nomenclature. 
Regnum Veg. 46. Utrecht. 

, & F. A. STAFLEU. 1964. Index Herbariorum. Part. 1. 
The herbaria of the world. 5th ed. Regnum Veg. 31. Utrecht. 

Lascut, A. J. M. 1889. Examination of some of the poisonous 
Ericaceae of North America. Pharm. Rund. 7: 208-213. 

LINNAEUS C. 1753. Species Plantarum. Stockholm. 

—— —————. 1754. Genera Plantarum. 5th ed. Stockholm. 

LirkE, R. 1968. Pflanzenreste aus dem untermiozán in Nordwest- 
sachsen. Palaeontographica Abt. B. 123: 173-183. 

LovELL, J. H. 1926. Honey plants of North America. Medina. 

, & H. B. LovELL. 1934. The pollination of Kalmia 
angustifolia. Rhodora 36: 25-28. 

Lovett, H. B. 1951. The mechanics of pollination. Bees 5: 18-23. 

Lupwic, C. F. 1783. Die neuere wilde baumzucht in einem alpha- 
betischen und systematischen verzeichisse. Leipzig. 

MACOUN, J. 1884. Catalogue of Canadian Plants. Part II. Gamo- 
petalae. Montreal. 

MansH, C. D., & A. B. CLAWSON. 1930. Mountain-laurel ( Kalmia 
latifolia) and sheep laurel (Kalmia angustifolia) as stock- 
poisoning plants. U. S. Dept. Agr. Tech. Bull. 219: 1-22. 

MATTHEWS, J. R., & E. M. KNox. 1926. The comparative morphology 
of the stamens in the Ericaceae. Trans. Proc. Bot. Soc. Edinb. 
29: 243-281. 

Matusow, H. 1897. Analysis of the root of Kalmia latifolia. Am. 
Jour. Pharm. 69: 341-343. 

MERRILL, E. D. 1945. In defense of the validity of William Bar- 
tram's binomials. Bartonia 23: 10-35. 

MEYER, B. S. 1927. The measurement of the rate of water-vapor 
loss from leaves under standard conditions. Am. Jour. Bot. 14: 

396 Rhodora [Vol. 76 

MILLER, C. C. 1895. Honey from mountain laurel. Am. Bee Jour. 
35: 825-826. 

MOUILLEFERT, P. 1897. Traite des Arbres & Arbrisse. Vol. II. 

MUENSCHER, W. C. 1957. Poisonous plants of the United States. 
Revised ed. New York. 

Munns, E. N. 1938. The distribution of important forest trees of 
the United States. U. S. Dept. Agr. Misc. Publ. 287: 1-176. 

NICHOLS, G. E. 1934. The influence of exposure to winter tempera- 
ture upon seed germination in various native American plants. 
Ecology 15: 364-373. 

NICHOLSON, G. 1896. Hand-list of trees and shrubs grown in Arbo- 
retum. Part II. Gamopetalae to Monocotyledons. Royal Gardens, 

NICHOLSON, S. M, & J. F. CrLovis. 1967. Dye plants and dye 
methods in West Virginia. Castanea 32: 111-116. 

NORTON, A. H. 1913. Some noteworthy plants from the islands and 
coast of Maine. Rhodora 15: 137-143. 

1935. Dr. Aaron Young, Jr. and the botanical survey 
of Maine. Rhodora 37: 1-16. 

PARKER, J. 1961. Seasonal changes in cold resistance of some north- 
eastern woody evergreens. Jour. Forest. 59: 108-111. 

PASCHKIS, H. 1881. Contributions to a closer knowledge of some 
little known leaves. Pharm. Jour. Trans. 12: 85-87. 

PELLETT, F. C. 1920. American honey plants. Hamilton. 

PENHALLOW, D. P. 1882. Fertilization of Kalmia. Gard. Month. 
24: 54. 

PERKINS, A. E. 1935. Notes on some rare plants of York County, 
Maine. Rhodora 37: 415-416. 

PETERSON, E. B. 1965. Inhibition of Black Spruce primary roots 
by a water-soluble substance in Kalmia angustifolia. Forest Sci. 
11: 473-479. 

PLUKENET, L. 1696. Opera omnia botanica in sex tomos divisa. 
IV. Almagestum botanicum. London. 

PORCHER, F. P. 1869. Resources of the Southern field and forests. 

PRITCHARD, W. R. 1956. Laurel (Kalmia angustifolia) poisoning 
of sheep. N. Am. Veterinarian 37: 461-462. 

PuRCHAS, S. 1625. Purchas His Pilgrimes. 3rd. Part. 16: 581-595. 

PunsH, F. 1814. Flora Americae Septentrionalis. London. 

RADFORD, A. E., H. E. AHLES, & C. R. BELL. 1968. Manual of the 
vascular flora of the Carolinas. Chapel Hill. 

RAFINESQUE, C. S. 1830. Medical Flora; or manual of the medical 
botany of the United States of North America. Vol. 2. Phila- 

1974] Genus Kalmia — Ebinger 397 

1840. Autikon Botanikum, Philadelphia. 

REHDER, A. 1910. Notes on the forms of Kalmia latifolia. Rhodora 
12: 1-3. 

. 1945. Notes on some cultivated trees and shrubs, II. 
Jour. Arnold Arb. 26: 472-481. 

1949. Bibliography of cultivated trees and shrubs 
hardy in the cooler temperate regions of the Northern Hemi- 
sphere. Jamaica Plain. 

REIFSNYDER, W. E. 1961. Seasonal variation in the moisture con- 
tent of the green leaves of Mountain Laurel. Forest Sci. 7: 

RoTHROCK, J. T. 1867. The fertilization of flowering plants. Am. 
Nat. 1: 64-72. 

RYDBERG, P. A. 1954. Flora of the Rocky Mountains and adjacent 
plains. New York. 

SARGENT, C. S. 1890. New or little known plants. A curious form 
of Kalmia. Garden Forest 3: 452. 

1893. Silva of North America, Vol. 3. Boston. 

Scott, G. D. 1962. Kalmia polifolia in Scotland. Glasgow Nat. 
18: 196-197. 

ScoTTER, G. W. 1966. A contributfon to the flora of the eastern 
arm of Great Slave Lake, Northwest Territories. Canad. Field- 
Nat. 80: 1-18. 

SHANKS, R. E. 1952. Notes on woody plant distribution in Ten- 
nessee. Castanea 17: 90-96. 

SMALL, J. K. 1903. Flora of the Southeastern United States. New 

1914. Ericaceae. North Amer. Flora 29: 33-102. 

SMITH, J. 1624. The general history of Virginia, New England, and 
the Summer Isles. Book 5. 

SMITH, W. W., A. R. HopGDoN, & R. EGGERT. 1947. Progress re- 
port on chemical weed control in blueberry fields. Proc. Am. 
Soc. Hort. Sci. 50: 233-238. 

SPRAGUE, E. 1871. The Rhododendron and “American Plants.” 

SPRENGEL. C. K. 1793. Das entdeckte Geheimnifs der Natur im 
Bau und in der Befruchrung der Blumen. Berlin. 

STANTEY. O. B. 1931. Fat deposits in certain Ericaceae. Butler 
Univ. Bot. Stud. 2: 33-41. 

STONE, G. E. 1909. A remarkable form of Kalmia latifolia. Rho- 
dora 11: 199-200. 

TALLENT. W. H., M. L. RIETHOF, & E. C. HORNING. 1957. Studies 
on the occurrence and structure of acetylandromedol (androme- 
dotoxin). Jour. Am. Chem. Soc. 79: 4548-4554. 

THIERET. J. W. 1963. Kalmia polifol’a: Second record from the 
Arctic. Canad. Field-Nat. 77: 173. 

398 Rhodora [Vol. 76 

TREVETT, M. F. 1956. Control of lambkill (Kalmia angustifolia) in 
lowbush blueberry fields. Northeastern Weed Control Conference 
Proc. 10: 193-198. 

TRUMPY, J. R. 1893. Propagating Kalmias. Gardening 1: 122. 

VAN RENSSELAER, M. G. 1889. The choice of a national flower. 
Garden Forest 2: 333-335. 

WAHLENBERG, W. G., & W. T. DOOLITTLE. 1950. Reclaiming Appa- 
lachian brush lands for economic forest production. Jour. Forest. 
48: 170-174. 

WANGENHEIM, F. U. J. 1788. Beschreibung der Polenblattrigen 
Kalmia und der gelbbluhenden Rosskastanie. Schr. Berlin Ges. 
Naturf. Freunde 8: 129-133. 

Waup, R. A. 1940. The action of Kalmia angustifolia (Lambkill). 
Jour. Pharm. Exp. Therap. 69: 103-111. 

Woop, C. E. JR. 1961. The genera of Ericaceae in the South- 
eastern United States. Jour. Arnold Arb. 42: 10-80. 

Woop, G. B., & F. BACHE. 1851. The Dispensatory of the United 
States of America. Philadelphia. 

1954. Andromedotoxin. A potent hypotensive agent from Rhodo- 
dendron maximum. Jour.“Am. Chem. Soc. 76: 5689-5692. 

Woop, T. F. 1883. Is sheep laurel poisonous to sheep? Am. Agr. 
42: 66. 



With increasing marine research activity in the Narra- 
gansett Bay area, the need for information about local 
marine algae has become urgent. Therefore, the available 
information has been compiled (Wood, in press) and a list 
of species known to occur in the State is presented. 

The earliest serious collecting of algae in Rhode Island 
seems to have been by S. T. Olney and his associates, and 
considerable effort was made in the years of 1846-1849. 
But, also, during the summer of 1846, J. W. Bailey visited 
Rhode Island and collected with Olney. Bailey, working 
independently, obtained specimens from Europe and worked 
up the collections systematically and included the Rhode 
Island records in his publication of 1847. Olney, on the 
other hand, sent his material to W. H. Harvey of Dublin, 
who identified them. The records were published by Olney 
in 1847, the same year that Bailey's appeared. Whose work 
appeared first is not clear, but Olney was the first Rhode 
Islander to publish on the marine algae of the State, while 
Bailey was probably the first worker to both study and 
publish on algae of Rhode Island. Subsequent records are 
those by Harvey (1852-1858), Olney (1871), Farlow (1873, 
1875, 1876, and 1881), and Bennett (1888), and the several 
reports by Collins between 1882 and 1896 which were 
largely summarized in his works of 1900 and 1909-1918. 
More recent reports are those by Wood & Palmatier (1954), 
the dissertation by Villalard (1971), and thesis by Vaughan 
(1971). In addition, three collections of concern are the 
*Algae Rhodiaceae" by Olney (1871) and the two large 
exsiccata sets in which Rhode Island algae occur — Farlow, 
Anderson & Eaton’s (1877-1889) “Algae Exsiccatae Ameri- 
cae-Borealis” and Collins, Holden & Setchell’s (1895-1919) 
“Phycotheca Boreali-Americana.” 

The senior author (rdw) prepared a list of species re- 
corded from Rhode Island by Taylor (1957), and then 


400 Rhodora [Vol. 76 

searched the literature for other early records. The sup- 
porting specimens were sought and examined, especially 
those at Brown University (courtesy of G. L. Church and 
W. H. Snell) and at the Farlow Herbarium at Harvard 
University (courtesy of I. M. Lamb). The complex of iden- 
tifications and synonyms required careful tracing, but veri- 
fication of most early names by examination of the actual 
collections was possible. The background information is to 
appear elsewhere (Wood, in press). The final list was 
checked by the junior author (mvb), and the latter also 
compiled the currently suggested synonyms that appear 
parenthetically in the list. Lastly, an evaluation of the 
frequency of occurrence of each taxon, based upon the 
authors’ personal observations, was included. Where identi- 
fications were still uncertain, Dr. Wm, Randolph Taylor 
graciously aided. The final manuscript was examined and 
edited by Urda K. Wood. 

It is hoped that this list will provide a background for 
future work on the algae of Rhode Island waters. 


The nomenclature below follows Taylor (1957), but other 
suggested names and references are indicated parentheti- 
cally. However, author citations have been checked, and 
those involving “in” and “ex” entries include both authors. 
Species reported for the first time in Rhode Island are desig- 
nated by an asterisk (*). The earliest known published rec- 
ord for each species in Rhode Island is cited in the second 
column, the abbreviations being: B — Bailey (1847), BENN. 
= Bennett (1888), B,o = Bailey (1847) and Olney (1847), 
c = Collins (1900), F = Farlow (1873), o = Olney (1847), 
OAR [for Olney’s “Algae Rhodiaceae"] = Olney (1871), PBA 
= in Collins, Holden & Setchell (1895-1919), T = Taylor 
(1937), (VA) = thesis by Vaughan (1971), (VI) = disserta- 
tion by Villalard (1971), W = Wood (1962), w&P = Wood 
& Palmatier (1954). 

Species which should be expected in the state, based on 

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 401 

Taylors (1957) range records, but for which no adequate 
report is yet available, are summarized at the end of the list. 

The frequency of occurrence of each species based on our 
combined personal observations is indicated in the third 
column as: t = trace, rare, + = local and uncommon, ++ 
= locally common, +++ = general and quite common. 
Where a rare taxon has been seen by only one of us, it 
is indicated as by “rdw” or “mvb” for R. D. Wood or M. 
Villalard-Bohnsack, respectively. 


1. Stichococcus marinus 
(Wille) Hazen T + 
2. Ulothrix flacca 
(Dillw.) Thuret in Le Jol. C dar 


3. Pseudendoclonium submarinum 
Wille T 


4. Gomontia polyrhiza 
(Lagerh.) Born. et Flah. T 

* 5. Capsosiphon fulvescens 
(C.Ag.) Setch. et Gard. (VI) * 
* 6. [Enteromorpha ahlneriana 
Blid.] (VI) t 
7. E. clathrata (Roth) J. Ag. OAR + 
(E. clathrata (Roth) Grev. — 
Parke & Dixon, 1964) 
8. E. compressa (L.) Grev. B,O + 
(ecotype of E. intestinalis (L.) 
Link — DaSilva & Burrows in 
Norton, 1972, p. 385) 
9. E. cruciata Collins OAR 




. E. erecta, (Lyngb.) J. Ag. 
(E. flexuosa ssp. paradoxa 
(Dillw.) Blid. — Bliding, 1963) 

* 11. E. groenlandica (J.Ag.) Setch. 









et Gard. 

E. intestinalis (L.) Link 

E. linza (L.) J. Ag. 

E. marginata J. Ag. 
(Blidingia marginata (J. Ag.) 
Dang. — Dangeard, 1958) 

E. minima Nag. 

(Blidingia minima (Nag. ex 
Kitz.) Kylin — Kylin, 1947) 
E. prolifera (Müll.) J. Ag. 
Monostroma fuscum (Post. et 
Rupr.) Witt. 

(Ulvaria obscura (Kütz.) Gayr. 
— Gayral, 1965) 

M. grevillei (Thuret) Witt. 
(Ulvopsis grevillei (Thuret) 
Gayr. — Gayral, 1965) 

. M. leptodermum Kjellm. 

(Kornmannia leptoderma 

(Kjellm.) Blid. — Bliding, 


M. oxyspermum (Kütz.) Doty 

(Ulvaria oxyspermum (Kiitz.) 

Blid. — Bliding, 1968) 

M. pulchrum Farlow 

. Percursaria percursa (C. Ag.) 

Ulva lactuca L. 
var. latissima (L.) DC. 

Prasiola stipitata Suhr. in Jess. 





[Vol. 76 



1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 403 


25. Chaetomorpha aerea (Dillw.) 
Kütz. B + 
(Ch. linum (Müll. Kütz. — 
Christensen, 1957) 
26. Ch. linum (Miill.) Kütz. B ++ 
27. Ch. melagonium (Web. et 
Mohr) Kiitz. t(rdw) 
28. Cladophora albida (Huds.) 
Kiitz. F tT 
29. Cl. flexuosa (Dillw.) Harv. C + 
(Cl. serica, (Huds.) Kütz. — 
Hoek, 1963) 
f. densa Collins PBA 
30. Cl. glaucescens (Harv.) Harv. F 
(Cl. albida (Huds.) Kutz. — 
Hoek, 1963) 
31. Cl. gracilis (Griff. ex Harv.) 
Kütz. [o rain 
(Cl. serica (Huds.) Kütz. — 
Hoek, 1963) 
32. Cl. hutchinsiae (Dillw.) Kütz. 
var. distans (C. Ag.) Kütz. T 
(Cl. hutchinsiae (Dillw.) Kütz. 
— Hoek, 1963) 
* 83. Cl. laetevirens (Dillw.) Kütz. (VI) m 
34. Cl. magdalenae Harv. F 
(Cl. aibida (Huds.) Kütz. — 
Hoek, 1963) 
35. Cl. refracta (Roth) Kütz. O 
(Cl. albida (Huds.) Kütz. — 
Hoek, 1963) 
36. Cl. rudolphiana (C. Ag.) Harv. C 
(Cl. serica (Huds.) Kütz. — 
Hoek, 1963) 
37. Cl. rupestris (L.) Kütz. OAR zu 
38. Rhizoclonium riparium (Roth) 
Harv. C t 


* 89. 


* A1. 

* 42. 






var. implexum (Dillw.) 

var. polyrhizum (Lyngb.) 

R. tortuosum Kütz. 

(R. riparium (Roth) Harv. — 
Scagel, 1966) 

Spongomorpha arcta (Dillw.) 

(Acrosiphonia arcta (Dillw.) 
J. Ag. — Kornmann, 1962) 
S. lanosa (Roth) Kütz. 

(S. aeruginosa (L.) Hoek — 
Hoek, 1963) 

S. spinescens Kütz. 
(Acrosiphonia arcta (Dillw.) 
J. Ag. — Kornmann, 1962) 
Urospora penicilliformis 
(Roth) Aresch. 


Bryopsis plumosa (Huds.) 
C. Ag. 

Codium fragile (Sur.) Hariot 
ssp. tomentosoides (van Goor) 



Ectocarpus confervoides 
(Roth) Le Jol. 

(E. siliculosus (Dillw.) 
Lyngb. — South & Cardinal, 

var. pygmaeus (Aresch.) 




[Vol. 76 





1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 405 








E. dasycarpus Kuck. 

E. fasciculatus Harv. 

E. penicillatus (C. Ag.) Kjellm. 
E. siliculosus (Dillw.) Lyngb. 
E. tomentosus (Huds.) Lyngb. 
(Spongonema tomentosum 
(Huds.) Kütz. — Kuckuck, 

Giffordia granulosa 

(Smith) Hamel 

G. mitchellae (Harv.) Hamel 
(G. mitchelliae (Harv.) Hamel 
— Earle, 1969) 

G. ovata (Kjellm.) Kylin 

G. secunda (Kiitz.) Batt. 
Pylaiella littoralis (L.) Kjellm. 
(Pilayella littoralis (L.) 
Kjellm. — Dixon & Russell, 

Sorocarpus micromorus 
(Bory) Silva 

Streblonema aecidioides 
(Rosenv.) Fosl. 

(Entonema aecidioides (Rosenv.) 
Kjellm. — Edelstein & 
McLachlan, 1968) 

. S. chordariae (Farl.) De Toni 


. Cladostephus verticillatus 

(Lightf.) Lyngb. 

. Sphacelaria cirrosa 

(Roth) C. Ag. 

. S. radicans (Dillw.) C. Ag. 


. Ascocyclus orbicularis Magnus 

(Myrionema magnusii (Sauv.) 
Lois. — Loiseaux, 1967) 



+ (rdw) 














Hecatonema terminalis 
(Kiitz.) Kylin 

Myrionema corunnae Sauv. 
M. foecundum (Stromf.) Fosl. 
(Hecatonema foecundum 
(Strómf.) Lois. — Parke & 
Dixon, 1968) 

M. globosum (Reinke) Fosl. 
(Microsporangium globosum 
Reinke — Parke & Dixon, 1964) 
M. strangulans Grev. 

Ralfsia borneti Kuck. 
R. clavata (Harv. in Hook.) 
Crouan frat. 
R. pusilla (Strómf.) Batt. 
R. verrucosa (Aresch.) J. Ag. 


(Elachistaceae — Parke & Dixon (1964) ) 

Elachistea chondri Aresch. 
(Elachista chondri Aresch. — 
Fritsch, 1945; Parke & Dixon, 

E. fucicola (Vell.) Aresch. 

( Elaschista fucicola Aresch. — 
Fritsch, 1945; Parke & Dixon, 

Halothrix lumbricalis 

(Kütz.) Reinke 

Chordaria flagelliformis 
(Müll.) C. Ag. 
var. densa Farl. 
Eudesme virescens (Carm. ex 
Harv. in Hook.) J. Ag. 


[Vol. 76 




1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 407 

78. Leathesia difformis (L.) 

Aresch. OAR ++ 
79. Sphaerotrichia divaricata 
(C. Ag.) Kylin OAR + 
80. Stilophora rhizodes 
(Turn.) J. Ag. OAR 
81. Arthrocladia villosa 
(Huds.) Duby C t 
82. Desmarestia aculeata (L.) 
Lamour. B + 
83. D. viridis (Müll. Lamour. B "n 
84. Striaria attenuata 
(Grev.) Grev. C 
* 85. Asperococcus echinatus 
(Mert.) Grev. (VI) Tom 

(A. fistulosus (Huds.) Hook. — 
Edelstein, Wynne & McLachlan, 

86. Desmotrichum undulatum 

(J. Ag.) Reinke B,O 
87. Litosiphon filiformis 

(Reinke) Batt. PBA 
88. Myriotrichia clavaeformis 

Harv. C 
89. M. filiformis Harv. F 
90. Petalonia fascia 

(Müll. Kuntze BENN. ++ 
91. Punctaria latifolia Grev. BENN. + 
92. P. plantaginea (Roth) Grev. O ule ce 
93. Rhadinocladia cylindrica 

Schuh T 

94. R. farlowii Schuh T 












Scytosiphon lomentaria 
(Lyngb.) Link 

(S. lomentarius (Lyngb.) Link 
— Farlow, 1881) 

Dictyosiphon chordaria 
Aresch. (as Gobia) 
D. foeniculaceus 
(Huds.) Grev. 


Alaria esculenta (L.) Grev. 
Chorda filum (L.) Lamour. 
C. tomentosa Lyngb. 
Laminaria agardhii Kjellm. 
(L. saccharina (L.) Lamour. — 
Wilce, 1965) 

L. digitata (Huds.) Lamour. 
L. intermedia Fosl. 

(L. digitata (Huds.) Lamour. 
— Sundene, 1958) 

L. longicruris De la Pyl. 

L. saccharina (L.) Lamour. 


Ascophyllum mackaii 
(Turn.) Holmes et Batt. 

(A. nodosum (L.) Le Jol. — 
Gibb, 1957) 

A. nodosum (L.) Le Jol. 
Fucus edentatus De la Pyl. 
(F. distichus L. ssp. edentatus 
(De la Pyl.) Pow. — Powell, 

F. evanescens C. Ag. 

(F. distichus L. ssp. 
evanescens (C. Ag.) Pow. — 
Powell, 1963) 



[Vol. 76 

++ +++ 



+ (rdw) 

++, +++ 


1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 409 

110. F. spiralis L. B res 
var. limitaneus Montagn. T 

111. F. vesiculosus L. B,O ro 

112. Sargassum filipendula C. Ag. B,O + 

*113. S. natans (L.) J. Meyen (VA) t(drift) 


114. Bangia ciliaris Carm. ex Harv. C t 

(Erythrotrichia ciliaris (Carm. 
ex Harv.) Thur. in Le Jol. — 
Parke & Dixon, 1964) 

115. B. fuscopurpurea 

(Dillw.) Lyngb. B,O FT 
116. Erythrotrichia carnea 
(Dillw.) J. Ag. C 
117. Goniotrichum alsidit 
(Zanard.) Howe C t 
118. Porphyra leucosticta 
Thuret in Le Jol. t 
*119. P. miniata (C. Ag.) C. Ag. (VI) x 
120. P. umbilicalis (L.) J. Ag. B,O dT 
121. Acrochaetium daviesti 
(Dillw.) Nag. C 
122. A. flexuosum Vickers T 
123. Kylinia secundatum 
(Lyngb.) Papenf. C 

(Acrochaetium secundatum 
(Lyngb.) Nig. — Parke & 
Dixon, 1968) 
124. K. virgatula (Harv.) Papenf. C 
(Acrochaetium virgatula 
(Harv.) J. Ag. — Parke & 
Dixon, 1968) 











Rhodochorton penicilliforme 
(Kjellm.) Rosenv. 

R. purpureum 

(Lightf.) Rosenv. 


Nemalion multifidum 
(Web. et Mohr) J. Ag. 

(N. helminthoides (Vell. in 
With.) Batt. — Parke & 
Dixon, 1964) 


Scinaia furcellata 
(Turn.) Bivona 

Asparagopsis hamifera 

(Hariot) Okam. (VI), (VA) 

(Bonnemaisonia hamifera Har. 
— Feldmann & Feldmann, 

. Gelidium crinale 
(Turn.) Lamour. 

Dumontia incrassata 
(Miill.) Lamour. 

Polyides caprinus 
(Gunn.) Papenf. 
(P. rotundus (Huds.) Grev. 
— Parke, 1953) 

Hildenbrandia prototypus 

[Vol. 76 


+ (mvb) 




1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 411 

134. Rhododermis georgii 
(Batt.) Collins T 
(Rhodophysema georgii Batt. 
—Dixon, 1964) 

135. Corallina officinalis L. BENN. mtm 
136. Fosliella farinosa 
(Lamour.) Howe e 

(Melobesia farinosa Lamour. 
— Hamel & Lemoine, 1952) 
137. F.lejolisii (Rosan.) Howe C + 
(Melobesia lejolisit Rosan. — 
Hamel & Lemoine, 1952) 
138. Lithophyllum corallinae 

(Crouan frat.) Heydr. PBA 
139. L. macrocarpum 
(Rosan.) Fosl. T 

(Dermatolithon pustulatum 
(Lamour.) Fos]. — Edelstein 
& McLachlan, 1968) 

140. L. pustulatum 

(Lamour.) Fosl. OAR 
*141. L. lenormandi 
(Aresch. in J. Ag.) Fosl. (VI) oe 

(Phymatolithon lenormandii 
(Aresch. in J. Ag.) Adey — 
Adey, 1966) 
142. Melobesia membranacea 
(Esper) Lamour. C 
(Epilithon membranaceum 
(Esper) Heydr. — Parke & 
Dixon, 1964) 
143. Phymatolithon laevigatum 
(Fosl.) Fosl. T ++ 
(Lithothamnion laevigatum 
Fos]. — Parke & Dixon, 1964) 











P. polymorphum (L.) Fosl. C 
(Lithothamnion polymorphum 

(L.) Aresch. in J. Ag. — Parke 

& Dixon, 1964) 

Gloiosiphonia capillaris 
(Huds.) Carm. ex Berk. C 
Euthora cristata 
(C. Ag.) J. Ag. F 
Choreocolax polysiphoniae 
Reinsch C 
Harveyella mirabilis 
(Reinsch) Reinke T 
Agardhiella tenera 
(J. Ag.) Schmitz B 

(as used by Taylor (1957), but 
designated A. baileyi (Harv. ex 
Kütz.) Taylor in Taylor & 
Rhyne (1970: 13), and since 
changed to Neoagardhiella 
baileyi (Harv. ex Kütz.) Wynne 
& Taylor, Wynne & Taylor, 

1973, p. 101) 
Cystoclonium purpureum 
(Huds.) Batt. B,O 
var. cirrhosum Harv. C 

Hypnea musciformis 

(Wulf.) Lamour. BENN. 

[Vol. 76 




++ +++ 

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 413 

152. Gracilaria foliifera 
(Forssk.) Borg. (0) t 
153. G. verrucosa (Huds.) Papenf. W&P +++ 
154. Ahnfeltia plicata 
(Huds.) Fries B,O T 
155. Gymnogongrus griffithsiae 
(Turn.) Mart. C t(mvb) 
156. Phyllophora brodiaei 
(Turn.) Endl. F + 

(P. truncata (Pallas) Newr. et 
Taylor — Newroth & Taylor, 

var. catenata (C. Ag.) Collins T 
157. P. membranifolia 
(Good. et Woodw.) J. Ag. 0 mo 

(P. pseudoceranoides (Gmel.) 
Newr. et Taylor — Newroth & 
Taylor, 1971) 

158. P. traillii Holmes ex Batt. C 
159. Chondrus crispus Stackh. B,O ttt 
160. Gigartina stellata 
(Stackh.) Batt. G M 

(Palmaria palmata (L.) 
Stackhouse — Guiry, 1974, 

p. 219.) 
161. Rhodymenia palmata 
(L.) Grev. B,O TARERE 
162. Champia parvula 
(C. Ag.) Harv. B,O bad a 

163. Lomentaria baileyana 
(Harv.) Farl. OAR TRT 















[L. clavellosa (Turn.) Gail.] 
L. orcadensis 
(Harv.) Coll. ex Taylor 


Antithamnion americanum 
(Harv.) Farl. in Kjellm. 

A. cruciatum (C. Ag.) Nag. 
A. plumula 

(Ellis) Thuret in Le Jol. 

A. pylaisaei (Mont.) Kjellm. 
Callithamnion baileyi Harv. 
var. laxum Farlow 

C. byssoides 

Arnott ex Harv. in Hook. 

C. corymbosum 

(Smith) Lyngb. 

C. roseum (Roth) Lyngb. 

C. tetragonum (With.) C. Ag. 
Ceramium deslongchampii 
Chauv. in Duby 

C. diaphanum (Lightf.) Roth 
C. fastigiatum Harv. 

C. rubriforme Kylin (prox.) 
C. rubrum (Huds.) C. Ag. 
var. pedicellatum Duby 

(C. pedicellatum DC. — Parke 
& Dixon, 1964) 

var. squarrosum Harv. 

C. strictum Harv. 

Griffithsia globulifera Harv. 
G. tenuis C. Ag. 
Pleonosporium borreri 
(Smith) Näg. ex Hauck 
Plumaria elegans 

(Bonnem.) Schmitz 

Ptilota serrata Kütz. 

[Vol. 76 

+ (mvb) 
BENN. + 
OAR t 
B =t- 
C + 
B,O +,+ + 
OAR ++ 
(VA) t? 
B,O +++ 
C + 
C + 
O0 t 
F Tte 
B?,F t 

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 415 










Seirospora griffithsiana Harv. 
(S. seirosperma (Harv.) 
Dixon — Dixon, 1964) 
Spermothamnion turneri 
(Mert.) Aresch. 

(S. repens (Dillw.) Rosenv. — 
Rosenvinge, 1924) 

Spyridia filamentosa 

(Wulf.) Harv. in Hook. 

T railliella, intricata 

(J. Ag.) Batt. 

(sporophyte of Asparagopsis 
hamifera — Taylor, 1957, 

p. 291) 

Grinnellia americana 
(C. Ag.) Harv. 
Phycodrys rubens (L.) Batt. 


. Dasya pedicellata 
(C. Ag.) C. Ag. 
(D. baillouviana (Gmel.) Mont. 
— Dixon & Irvine, 1970) 

Chondria baileyana 
(Mont.) Harv. 
C. dasyphylla (Woodw.) C. Ag. 
C. sedifolia Harv. 
C. tenuissima 
(Good. et Wooaw.) C. Ag. 
Polysiphonia denudata 
(Dillw.) Grev. ex Harv. in 
. P. elongata Harv. in Hook. 
. P. fibrillosa (Dillw.) Spreng. 







416 Rhodora [Vol. 76 

200. P. harveyi Bail. 0 tt 
var. olneyi (Harv.) Collins O 
201. P. lanosa (L.) Tandy B,O ++, +++ 
202. P. nigra (Huds.) Batt. F + 
203. P. nigrescens (Huds.) Grev. B + 
204. P. novae-angliae Taylor B 
205. P. subtilissima Mont. F + 
206. P. urceolata 
(Lightf. ex Dillw.) Grev. C ++ 
207. Rhodomela confervoides 
(Huds.) Silva B,O +++ 

208. R. virgata Kjellm. C 


Codiolum gregarium A. Br. 
Ulothrix implexa Kütz. 

Bolbocoleon piliferum Pringsh. 
Entocladia flustrae (Reinke) Batt. 
(Epicladia flustrae Reinke — Parke & Dixon, 1968) 
E. wittrockii Wille 
(Ectochaete wittrockii (Wille) Kylin — Parke & Dixon, 1968) 
Pringsheimiella scutata (Reinke) Marchew. 
Protoderma marinum Reinke 
(Pseudendoclonium marinum (Reinke) Aleem et Schulz — Loiseaux, 


Enteromorpha plumosa Kütz. 
(E. flexuosa ssp. paradoxa (Dillw.) Blid. — Bliding, 1963) 

Chaetomorpha atrovirens Taylor 
Cladophora crystallina (Roth) Kütz. 
(Cl. vagabunda (L.) Hoek; Cl. glomerata (L.) Kütz. — Hoek, 1963) 
Cl. expansa (Mert.) Kütz. 
(Cl. vagabunda (L.) Hoek — Hoek, 1963) 

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 
Cl. flavescens (Roth) Kütz. 
(a doubtful name — Hoek, 1963) 

Derbesia vaucheriaeformis (Harv.) J. Ag. 

Bryopsis hypnoides Lamour. 

Ostreobium quekettii Born. et Flah. 
Vaucheria litorea C. Ag. 
V. thuretii Woronin 

Ectocarpus subcorymbosus Farl. 
(Farlowiella onusta (Kütz.) Kuck. — Kuckuck, 1956) 

(Elachistaceae — Parke & Dixon, 1964) 

Myriactula minor (Farl. Taylor 

Acrothrix novae-angliae Taylor 
Eudesme zosterae (Lyngb.) Kylin 
(Cladosiphon zosterae (J. Ag.) Kylin — Parke, 1953) 
Stictyosiphon subsimplex Holden 

Desmotrichum balticum Kiitz. 
(D. undulatum (J. Ag.) Reinke — Kylin, 1947a) 
Sargassum fluitans Borg. 

Asterocytis ramosa (Thwaites in Harv.) Gobi ex Schmitz 

Audouinella membranacea (Magnus) Papenf. 


418 Rhodora [Vol. 76 


Lithothamnium laeve (Strómf.) Fosl. 
(Leptophytum laeve (Strómf.) Adey — Adey, 1966) 

Callithamnion circinatum (Kütz.) J. Ag. 

Membranoptera denticulata (Mont.) Kylin 


Bostrychia rivularis Harv. 
Polysiphonia flexicaulis (Harv.) Collins 


ADEY, W. H. 1966. The genera Lithothamnion, Leptophytum (nov. 
gen.) and Phymatolithon in the Gulf of Maine. Hydrobiologia 
28(3-4): 321-370. 

BAILEY, J. W. 1847. Notes on the algae of the United States, Am. 
J. Sci. & Arts 2: 80-85, 399-403. 

BENNETT, J. L. 1888. Plants of Rhode Island, being an enumeration 
of plants growing without cultivation in the State of Rhode 
Island. Published by the author, Providence Press Co., Provi- 
dence. 128 pp. 

BLIDING, C. 1963. A critical survey of European taxa in Ulvales, 
I, Capsosiphon, Percursaria, Blidingia, Enteromorpha. Opera 
Bot. 8(3): 1-160. 

1968. A critical survey of European taxa in Ulvales, 
II, Ulva, Ulvaria, Monostroma, Kornmannia, Bot. Not. 121: 

CHRISTENSEN, T. 1957. Chaetomorpha linum in the attached state. 
Bot. Tidsskr. 53: 311-316. 

CoLLINS, F. S. 1900. Preliminary lists of New England plants, 
V. Marine algae. Rhodora 2: 41-52. 

1909-1918. The green algae of North America. Tufts 
Coll. St. 2(3): 79-480; First Suppl, ibid. 3(2): 69-109, 1912; 
Second Suppl., ibid. 4(7): 1-106, 1918. 

CoLLINS, F. S., I. HOLDEN, & W. A. SETCHELL. 1895-1919. Phyco- 
theca Boreali-Americana. (Exsiccata) Fasc. 1-46 + A-E. 

DANGEARD, P. 1958. La reproduction et le développement de 
l'Enteromorpha marginata J. Ag. et le rattachement de cette 
espéce au genre Blidingia. C. R. Acad. Sci., Paris 246: 347-351. 

DixoN, P. S. 1964. Taxonomic and nomenclatural notes on the 
Florideae IV. Bot. Not. 117: 56-78. 

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 419 

Drxon, P. S., & L. M. IRVINE. 1970. Miscellaneous notes on algal 
taxonomy and nomenclature, III. Bot. Not. 123(4): 474-487. 
DixoN, P. S, & G. RUSSELL. 1964. Miscellaneous notes on algal 
taxonomy and nomenclature, I. Bot. Not. 117(3): 279-284. 
EARLE, S. A. 1969. Phaeophyta of the eastern Gulf of Mexico. 

Phycologia 7(2): 71-254. 

EDELSTEIN, T., & J. MCLACHLAN. 1968. Investigations of the marine 
algae of Nova Scotia, V. Additional species new or rare to 
Nova Scotia. Can. J. Bot. 46: 993-1003. 

EDELSTEIN, T., M. J. WYNNE, & J. MCLACHLAN. 1970. Melanosiphon 
intestinalis (Saund.) Wynne, a new record for the Atlantic. 
Phycologia 9(1): 5-9. 

Fartow, W. G. 1873. List of seaweeds or marine algae of the 
south coast of New England. Rep. U. S. Comm. Fish & Fisheries 
1: 281-294, 

1875. List of the marine algae of the United States 
with notes of new or imperfectly known species. Proc. Am. 
Acad. Arts & Sci. 10: 351-380. 

. 1876. List of the marine algae of the United States. 
Rep. U. S. Comm. Fish & Fisheries 5: 691-718. 

1881. The marine algae of New England. Rep. U. S. 
Comm. Fish & Fisheries for 1879, Append. A-1: 1-210. 

FARLOW, W. G., C. L. ANDERSON, & D. C. EATON. 1877-1889. Algae 
Exsiccatae Americae-Borealis. (Exsiccata) Fasc. 1-5. Boston. 

FELDMANN, J., & G. FELDMANN. 1942. Recherches sur les Bonne- 
maisoniacées et leur alternance de générations. Ann. Sci. Nat. 
Bot., Ser. II, 3: 75-175. 

FRITSCH, F. E. 1945. The structure and reproduction of the algae. 
Vol. II. The Univ. Press, Cambridge. 939 pp. 

GAYRAL, P. 1965. Monostroma  Thuret, Ulvaria Rupr., emend. 
Gayral, Ulvopsis Gayral. (Chlorophycées, Ulotrichales): struc- 
ture, reproduction, cycles, position systematique. Rev. Gén. Bot. 
72 (857): 627-638. 

GIBB, D. C. 1957. The free-living forms of Ascophyllum nodosum 
(L.) Le Jol. J. Ecol. 45(1): 49-83. 

Guiry, M. D. 1974. A reappraisal of the genus Palmaria Stack- 
house. Brit. Phyc. J. 9(2): 219. : 
HAMEL, G., & P. LEMOINE. 1952. Corallinacées de France et 
d'Afrique du Nord. Arch. Mus. Natn. Hist. Nat., Paris 7(1): 


Harvey, W. H. 1852-1858. Nereis boreali-americana. Part I. 
Smithsonian Contrib. Knowledge 3(4): 1-150; Part II, ibid., 
5(5): 1-258, 1853; Part III, tbid., 10: 1-140, 1858 (including 

HokK, C. VAN DEN. 1963. Revision of the European species of 
Cladophora. E. J. Brill, Leiden. 248 pp. 

420 Rhodora [Vol. 76 

KORNMANN, P. 1962. Eine Revision der Gattung Acrosiphonia. 
Helgolànd. Wiss. Meeres. 8: 219-242. 

Kuckuck, P. 1956. Ectocarpaceen-Studien IV. Herponema, Kutz- 
ingiella nov. gen., Farlowiella nov. gen. herausgegeben von Peter 
Kornmann. Helgolànd. Wiss. Meeres. 5: 292-325. 

1960. Eectocarpaceen-Studien VI. Spongonema. Helgo- 
lànd. Wiss. Meeres. 7: 93-113. 

KvLIN, H. 1947. Ueber die Fortpflanzungsverhaltnisse in der Ord- 

nung Ulvales. K. Fys. Sallsk. Lund Fórh. 17: 174-182. 
1947a. Die Phaeophyceen der Schwedischen Westküste. 
Lunds Univ. Arsskr. N. F., Avd. 2, 43(4) : 1-99. 

LorsEAUX, S. 1967. Morphologie et cytologie des Myrionémacées. 
Critères taxonomiques. Rev. Gén. Bot. 74(878): 329-947. 

Lunp, S. 1959. The marine algae of East Greenland, I. Taxo- 
nomical part. Medd. Gronland 156(1): 1-248. 

NEwRoTH, P. R., & A. R. A. TAYLOR. 1971. The nomenclature of 
the North American species of Phyllophora Greville. Phycologia 
10(1): 93-97. 

Norton, T. A. 1972. The marine algae of Lewis and Harris in the 
Outer Hebrides. British Phycol. J. 7(3): 375-385, 

OLNEY, S. T. 1847. Rhode Island plants, 1846, or additions to the 
published lists of the Providence Franklin Society. Proc. Provi- 
dence Franklin Soc. 1(2): 25-42. 

1871. Algae Rhodiaceae. A list of Rhode Island algae, 
collected and prepared by Stephen T. Olney, in the years 1846- 
1849, now distributed from his own herbarium. Published by 
the author, printed by Hammond, Angell & Co., Providence. 
13 pp. (Exsiccata, few sets). 

PARKE, M. 1953. A preliminary check-list of British marine algae. 
J. Mar. Biol. Ass. U. K. 32: 497-520. 

PARKE, M., & P. S. Dixon. 1964. A revised check-list of British 
marine algae. J. Mar. Biol. Ass. U. K. 44: 499-542. 

1968. Check-list of British marine algae — second re- 
vision. J. Mar. Biol. Ass. U. K. 48: 783-832. 

PowELL, H. T. 1963. Speciation in the genus Fucus L. and related 
genera. In: J. P. Harding & N. Tebble (ed.), Speciation in 
the sea. Syst. Ass. Publ. 5: 63-77. 

RosENvINGE, K. L. 1924. The marine algae of Denmark. Vol. III. 
Rhodophyceae. III. (Ceramiales). K. Danske Vidensk. Selsk., 
Ser. Nat. 7: 287-486. 

ScaGeL, R. F. 1966. Marine algae of British Columbia and northern 
Washington. Part I: Chlorophyceae (green algae). Nat. Mus. 
Can., Biol. Ser. No. 74, Bull. 207: 1-257. 

SOUTH, R., & A. CARDINAL. 1970. A check list of marine algae of 
eastern Canada. Can. J. Bot. 48: 2077-2095. 

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 421 

SUNDENE, O. 1958. Interfertility between forms of Laminaria digi- 
tata. Nytt. Mag. Bot. 6: 121-128. 

TAYLOR, W. R. 1937. Marine algae of the northeastern coast of 
North America. Univ. Michigan Press, Ann Arbor. 427 pp. 

1957. Ibid., 2nd ed. 509 pp. 

TAYLOR, W. R., & C. F. RHYNE. 1970. Marine algae of Dominica. 
Smithsonian Contrib. Botany 3: 1-16. 

VAUGHAN, C. F. 1971. An ecological survey of the macroscopic 
algae of Point Judith Pond (Salt Pond), Rhode Island. M. S. 
Thesis, University of Rhode Island. 179 pp. 

VILLALARD, M. 1971. Phytosociology of intertidal algae of Conani- 
cut Island, Rhode Island. Ph.D. Thesis. University of Rhode 
Island. 227 pp. 

WiLCE, R. T. 1965. Studies in the genus Laminaria. III. A re- 
vision of the North Atlantic species of the Simplices section of 
Laminaria. Bot. Goteborg. 3: 247-256. 

Woop, R. D. 1962. Codium is carried to Cape Cod. Bull. Torrey 
Bot. Club 89: 178-180. 

————— ————. (in press). Information for a list of marine algae of 
Rhode Island. New England Marine Resources Information Pro- 
gram, University of Rhode Island, Kingston. 

Woop, R. D, & E. A. PALMATIER. 1954. Macroscopic algae of the 
coastal ponds of Rhode Island. Am. J. Bot. 41(2): 135-142. 

WYNNE, M. J., & W. R. TAYLOR. 1973. The status of Agardhiella 
tenera and Agardhiella baileyi (Rhodophyta, Gigartinales). 
Hydrobiologia 43(15): 93-107. 






B. J. Cox! 

Systematics of the genus Lupinus is often viewed as one 
of the most perplexing studies ever attempted. This is 
attested by the diverse taxonomic treatments of various 
floras and special publications. While several students of 
the lupines have recognized many entities, others have 
found these plants in such confusing mosaics that enormous 
synonymic reductions were made; neither approach has 
solved the taxonomic problems formed by lupine diversity. 

In a biosystematic revision of the Lupinus lepidus-L. 
caespitosus complex of the Sierra Nevada, Cascade and 
Rocky Mountains, various techniques of study were em- 
ployed in an effort to show the degree of relationship be- 
tween the taxa (Cox 1972a). Over 75 epithets have been 
applied to various members of this complex since 1828 
when L. lepidus was described, and these epithets have 
been considered by some as referable to a single species. 

In order to present the concept of the L. lepidus-L. caes- 
pitosus complex in a digestible and economically feasible 
form, a taxonomic treatment of only Lupinus lyallii, its five 
subspecies and six varieties will be offered here. More 
complete distribution maps, cytological and chemotaxonomic 
data, photographs of typical specimens, and illustrations of 
typical leaflet and floral parts are given elsewhere (Cox 
1972a, 1972b, 1973a, 1973b). 

Chromosome numbers: Taxa within Lupinus lyallii have 
chromosome number reports of 2n — 48 (subsp. lyallii, var. 
lyallii and var. villosus, and subsp. minutifolius) by Cox 
(1972b). The chromosomes are extremely small and counts 
are difficult to obtain, undoubtedly explaining the lack of 
previous reports. Although Phillips (1957) reported a 

'Studies conducted at the Division of Biological Sciences, University 
of Missouri, Columbia, Missouri. 


1974] Genus Lupinus — Cox 423 

chromosome count for L. lyallii of n = 24, his study plant 
was L. minimus. Chromosome counts have offered little in 
determining the genetic affinities within the L. lepidus-L. 
caespitosus complex since all reported numbers are 2n — 48 
(Cox 1972b). 

Chemosystematic data. Seed proteins: Comparisons were 
made of seed proteins of four of the L. lyallii taxa by poly- 
acrylamide gel electrophoresis (Cox 1972a, 1973b). Con- 
siderable variation was found in several polymorphic popu- 
lations; largely, the electrophoresis disc protein data closely 
resemble those data from the morphological study in the 
degree of variation found. Although no protein band oc- 
curred in common with all taxa studied, likewise no pro- 
tein band was found unique to any one taxon. Each popu- 
lation was found to have a distinct combination of proteins, 
resulting in low percent similarity values between the taxa. 

Seed Alkaloids: The seed alkaloids of all of the sub- 
species and varieties of Lupinus lyallii were compared 
by thin-layer chromatography (TLC) separation by Cox 
(1973a). Although some interpopulational variation was 
found, the alkaloids were generally quite consistent. Corre- 
lations were made between alkaloid shapes, R: values, posi- 
tive color reactions with two different visualizing agents, 
and comparisons with standards. Percent similarity values 
were high among the interspecific taxa, and identical alka- 
loid combinations were shared among a few, these differing 
only in relative amounts of individual alkaloids. 

Morphological and. distributional considerations: Field 
and herbarium-laboratory studies were conducted over a 
three year period on the Lupinus lepidus-L. caespitosus 
complex. Techniques and preparations of specimens for 
measurements in the descriptions are given by Cox (1972a). 

The following herbaria are acknowledged for the use of 
over 1000 specimens of the taxa of Lupinus lyallii: AEC, 

424 Rhodora [Vol. 76 

Lupinus lyallii A, Gray 

Plants perennial, herbaceous, caespitose or mat-like, 
less than 18 cm. tall; leaflets few (5-9), oblanceolate, the 
largest less than 2 cm. long; banners glabrous, usually re- 
flexed below the midpoint. 

Distribution: The taxa are primarily found in the high 
Sierra Nevada and Cascade Mountains with a few popula- 
tions radiating out into isolated localities (Fig. 1). 


A. Plants with spreading villous-hirsute to shaggy pubes- 
CENCE. aooaa lee. B. 

B. Plants lees than 10 cm. tall; flowers 6-7 mm. long; 
largest leaflets 4-6 mm. long; Sierra Nevada, alpine, 

dwarf. ...... 1b. L. lyallii subsp. lyallii var. danaus. 
B. Plants usually taller; flowers over 7 mm. long; larg- 
est leaflets over 6 mm. long. ............ . C. 

C. Plants decumbent, mat-like; leaflets 6- T, largest 
10-13 mm. long; Sierra Nevada, with a few sta- 
tions in Klamath Co., Ore. 

. lf. L. lyallii subsp. lyallii var. villosus. 

C. Plants erect to semi-decumbent, branching, 10-18 
cm. tall; leaflets 5-6, largest 9-21 mm. long; 
Washoe Co., Nevada. ...... , 

ZEE . 9. L. lyallii subsp. washoensis. 
A. Plants with appressed to subappressed pubescence. .. D. 
D. Plants erect in densely branching clumps; 12-18 cm. 

tall; Rogue River-Crater Lake region, Oregon. .. E. 

E. Branching stems mostly woody, less than 1.5 mm. 
in diameter; leaflets 5-6; racemes 1.3-1.4 cm. in 
diameter ; flowers 6-7.5 mm. long. 

. 1c. L. lyallii subsp. lyallii var. fructiculosus. 

E. Branching stems mostly herbaceous, more robust; 
leaflets 7-9; racemes 1.9-2.5 cm. in diameter; 
flowers 7-9.5 mm. long. ................. s. 
e le. L. lyallii subsp. lyallii var. roguensis. 

D. Plants decumbent or erect and densely caespitose. 

1974] Genus Lupinus — Cox 425 

F. Plants mat-like, usually less than 10 cm. tall; 
flowers 6-7 mm. long; largest leaflets 4-6 mm. 
long; alpine, Sierra Nevada Range. .........- 
Mero Rd 1b. L. lyallii subsp. lyallii var. danaus. 

F. Plants usually larger or flowers longer. ..... G. 
G. Plants 4-6 cm. tall; flowers 7.5-10 mm. long; 

largest leaflets 6-9 mm. long, upper surface 
glabrous; Steens Mts, Harney Co., Oregon. 
TM IO ay e «= 3. L. lyallii subsp. minutifolius. 
G. Plants mostly taller; flowers various; largest 
leaflets various lengths, mostly pubescent 
(becoming somewhat glabrate in subsp. alcis- 
temporis). occ ccc ecw cece gien p hh n H. 
H. Plants mostly decumbent; Clearwater Co., 
Idaho. .. 2. L. lyallii subsp. alcis-temporis. 

H. Plants erect or mat-like, not decumbent. 
I Leaves all basal, erect, in small tufts; 
petioles 3-9 em. long; largest leaflets 

9-22 mm. long; SW Idaho & NE Ne- 
vada. .. 4. L. lyallii subsp. subpandens. 

I. Leaves cauline, basal, or both, generally 
ascending or more or less spreading, 
general not in small tufts; petioles 
usually 2-5.5 em. long; largest leaflets 

J. Flowers 10-12.5 mm. long; racemes 
2.5-4.5 (5.5) cm. long, 2.5-3.0 cm. 

in diameter ; leaflets 6-9; high alpine 

areas of western Cascade Range, 

Öre & Wash UU b e d nih s 1d. L. 

lyallii subsp. lyallii var. macroflorus. 

J. Flowers 7.5-10.5 mm. long; racemes 

1.5-3 em. long, 2.0-2.3 cm. in di- 
ameter; leaflets 5-7; alpine and sub- 

alpine, wide distribution, Sierra Ne- 

vada Range, Calif. & Cascade Range, 

426 Rhodora [Vol. 76 

Ore., Wash. & B. C. .......... la. 
L. lyallii subsp. lyallii var. lyallii. 

la. Lupinus lyallii subsp. lyallii var. lyallii 

Lupinus lyallii A. Gray, Proc. Amer. Acad. 7: 334. 
1868. Type: “Summit of the Cascade Mountains, lat 49°, 
elev. 7500 ft.”, Lyall, Jul 1860 (Holotype: K; Isotypes: 
BM, GH, POM). Photos: (RSA, UC, UMO, US). 

L. lepidus subsp. lyallii (Gray) Detl., Amer. Mid. Nat. 
45: 490-491. 1951. 

L. alpinus Heller, Muhlenbergia 6: 22. 1910. Type: 
NEVADA: WASHOE CO.: Mt Rose, Heller 9914a (Holo- 
type: Nev. Agr.; Isotypes: GH, ISC, MIN, NMC, NY, POM, 
RSA, UC, US, WS). Photos: (RSA, UMO). 

L. paulimus Greene, Leafl. Bot. Obs. & Crit. 2: 234. 
1912. Type: OREGON: DESCHUTES CO.: Paulina Lake, 
W end, Leiberg 550 (Holotype: US; Isotypes: GH, UC). 

L. rubro-soli Eastw., Leafl. West. Bot. 3: 203. 1941. 
Type: CALIFORNIA: MENDOCINO CO.: Red Mt., Eastwood 
& Howell 4674 (Holotype: CAS; Isotype: US). Photos: 
(RSA, UMO). 

Plants perennial, caespitose, forming mat-like radicle 
clumps, 15-25 cm. in diameter, from a thick, woody caudex 
which often branches in older plants, 6-17 cm. tall; foliage 
2-7 cm. tall; stems poorly developed, less than 1.5 mm. in 
diameter, semi-prostrate, some sprawling-decumbent, some 
erect, subappressed silky sericeous throughout, hairs usually 
less than 1 mm. long; cauline nodes 0-2; petioles slender, 
2-5(7.5) cm. long; stipules generally inconspicuous; leaf- 
lets 5-7, oblanceolate-linear to elliptic-obovate, apex acute to 
short mucronate, largest 5-12(18) mm. long, av. 9.95 mm., 
1.5-4.5 mm. wide, length to width ratio 2.17-4.0(6.0), av. 
3.18, appressed silky-sericeous on both surfaces; peduncles 
usually much exceeding the foliage, 3-8(10) cm. long; 
racemes capitate to subcapitate, 1.5-3 em. long, occasionally 
to 4.0 cm. at anthesis, 20-23 mm. in diameter, rarely 2X 
as long as wide; flowers dense, blue, 7.5-10.5 mm. long, av. 
9.44; bracts persistent to caducous, 3-6.5 mm. long, subu- 

1974] Genus Lupinus — Cox 427 

late to lance-linear; pedicels 1.5-2.5 mm. long; calyx bi- 
labiate, the upper-lip 3.5-6.0 mm. long, bilobed, the notch 
1-2.5 mm. deep, the lower-lip 3.8-6.8 mm. long, apex serrate 
to tridentate, the center tooth to 0.5 mm. long, lips connate 
1-2 mm., bracteoles usually present but inconspicuous, to 
0.5 mm. long, attached at the sinus, sericeous; banner ellip- 
tic-oblong to suborbicular, glabrous, 7-10 mm. long, 5.1-8.0 
mm. wide, length to width ratio 1.10-1.57, av. 1.29, reflexed 
below or near the midpoint, the reflexed to appressed ratio 
1-1.3, av. 1.14, angle 120-150°; wings 6.5-9.8 mm. long, 3.5- 
5.5 mm. wide, length to width ratio 1.56-2.25 av. 1.90, claws 
1.0-2.2 mm. long, glabrous; keel 2.5-3.0 mm. wide at the 
widest point, angle 105-111°, ciliated along the upper mar- 
gin near the acumen; ovules 3-5; pods 10-15 mm. long, 4-5 
mm. wide, short sericeous; seeds 2-4, 1.8-3.0 mm. X 2.6-3.5 
mm., 1-1.5 mm. thick, tan, beige, brown and green mottled. 

This variety is primarily alpine to subalpine, ranging 
latitudinally in its distribution from the high Sierra Nevada 
Mountains of California through the Cascade Range of 
Oregon and Washington into British Columbia. These 
plants may be found in several soil types, such as volcanic 
ash, disintegrated granite, sandy, gravelly rock of several 
compositions to shale. They usually occur on slopes associ- 
ated most frequently with Pinus contorta, P. murrayana, 
P. albicaulis and Tsuga mertensiana.  Altitudinally, the 
plants are found above timberline at 3660 m. down to about 
2439 m., or in the more northerly latitudes to about 1830 m. 
Flowering occurs between June and September; the plants 
flowering during the latter part of the season usually are 
found in sites of lingering snowbanks. 

Representative specimens: CANADA: British Columbia: Bomford 
Mt, Ashnola Dist, Barr 9351 (UBC); Chileotin, Copley 472 (v); 
Lillooet, Anderson, Jul 1916 (v); Manning Park, Hardy, Aug 1945 
(v); Mt McLean, Anderson 7987 (v); Mt Moor, Bird 3436 (UBC) ; 
Nicoamen River head, Dawson 51833 (CAN); Noaxe Lake, Brink, Aug 
1957 (UBC); Yalakom River, Sugden 118 (v). USA: California: 
ALPINE CO.: Carson Pass, Yates 3980 (UC). EL DORADO CO.: Desolation 
Valley, Abrams 12740 (GH, NY, POM, UC). FRESNO CO.: Humphreys 

428 Rhodora [Vol. 76 

Basin, headwaters Piute Creek, Quibell 5118 (UC). LASSEN CO.: 
Lassen Volcanic Natl. Park, Rickett, Aug 1968 (UMO). MADERA CO.: 
Mt Lyell, at Moraine Meadows, Akey 318 (UC). MARIPOSA CO.: Mt 
Connes, Tioga Pass, Ramsey & Ramsey 2496 (POM). MODOC CO.: 
Antelope Valley, Applegate 847 (US). MONO CO.: Bodie, 2.2 mi 
N-NW, Graham 274 (UC). NEVADA CO.: Mt Stanford, Sonne 54 
(MO). PLACER CO.: Snow Mt, Bolt 103 (UC). SHASTA CO.: Bolam 
Road, near the S.P. RR, Cooke 15003 (uc). SISKIYOU CO.: Bolam 
Creek Canyon, opposite Coquette Falls, Mt Shasta. Cooke 25691 
(UC). TEHAMA CO0.: King Creek Meadow, Lassen Volcanic Natl. 
Park, Heller 15426 (Mo, NY). TRINITY CO.: Fisher Ridge, E of 
Black Rock, North Yolla Bolly Mts, Munz 16743 (RSA). TULARE CO.: 
Golden Trout Creek, Hall & Hall 8417 (uc). TUOLUMNE CO.: Bond 
Pass, Winkelman 276 (Uc). Nevada: WASHOE CO.: Mt Rose, Ken- 
nedy 1148 (NY, UC, US). Oregon: CURRY CO.: Bear Camp Ranger 
Station, Leach 1692 (ORE). DESCHUTES CO.: Bend, Farnham & Beach 
97 (OSC). DOUGLAS CO.: Bailey Mt, Applegate 4121 (uc). JACKSON 
co.: Ashland Peak. Thompson 12383 (MO, NY, UC). JFEFFFPSON CO.: 
Hoodoo, Big Lake Turnoff, US Hwy 20, Isely & Isely 11299 (ISC, 
UMO). JOSEPHINE CO.: Free and Easy Trail, head of Babyfoot, 
Leach 2661 (ORE). KLAMATH CO.: btn Fort Klamath & Summit, 
Furlong et al, 1901 (UC). CRATER LAKE NATL. PARK: Summit Lake, 
Henderson 9634 (ws). LAKE CO.: Drews Valley. Austin 1649 (Us). 
LANE CO.: McKinzie Pass, Peck 5467 (WILLU). LINN CO.: Bie Lake, 
Hawley & Anderson, Jul 1927 (osc). WALLOWA CO.: Wallowa Mts, 
Aneroid, 2 mi S, Peck 17985 (uc). Washington: CHELAN CO.: Blewett 
Pass region, Grant, Aug 1929 (UC). CLALLAM co: Hurricane Ridge, 
Thompson 14226 (GH, NY, US). JEFFERSON CO.: Tron Mt, Thompson 
11049 (MO. NY, POM. US). OKANOGAN CO.: Bunker Hill summit, 
Edwards 32 (ws). PIERCE co.: Mt Rainier Natl. Park: Flett’s 
Glacier, Flett 3124 (ws). wHATCOM co.: Mt Baker. Heliotrope 
Ridge, Muenscher 8201 (GH, UC. US. WS). YAKIMA CO.: Hell Rearing 
Canyon rim, Langdon 116 (osc). 

The populations from the Wallowa Mountains (Wallowa 
Co., Oregon) are isolated from other Lupinus lyallii subsp. 
lyallii populations by 322 km.; the subsp. alcis-temporis 
occupies areas along the Clearwater and Selway Rivers in 
Idaho, but it is quite distinctive. The growth habit of the 
Wallowa populations appears to be typically that of var. 
lyallii specimens in the normal distribution, with short 
vegetation to 7 cm. tall, capitate inflorescences 2-2.5 cm. 
long, small flowers 6-8 mm. long and short leaves 7-10 mm. 
long. But, these plants hold in common with L. minimus a 

1974] Genus Lupinus — Cox 429 

strong woody branching caudex at above ground level, many 
stems in a clump with leaves appearing basal, not more 
than 2-3 cauline leaves, and silky pubescence. 

The specimens of Lupinus lyallii subsp. lyallii var. lyallii 
in Deschutes Co., Oregon (Leiberg 550 — GH, UC, US), de- 
scribed as L. paulinus by Greene, are somewhat larger than 
other L. lyallii plants of the immediate region but closely 
resemble several populations found in California. The 
racemes of some plants range from 3-4 cm. to 7.5 cm. long 
and from 16 mm. to 22 mm, in diameter, with the leaflets 
11-15 mm. long, the upper surface of the leaflets sparsely 
pubescent, flowers approaching 9 mm. long and peduncles 
6-7 cm. long. Several plants resemble L. sellulus var. lobbii 
with the flowers scattered in verticils somewhat like L. 
sellulus var. medius (Cox 1973c). However, the population 
sample indicates that these variants are more closely 
related to L. lyallii subsp. lyallii var. lyallii than to any 
other taxon and not distinctive enough to warrant a sepa- 
rate epithet. 

The material found on Mt. Rose in Nevada, which Heller 
called Lupinus alpinus, flowers from late July to late Sep- 
tember and occurs at elevations of 2927-3110 m. 

The plant habit is low and spreading mats, much like 
that on Mt. Rainier in Washington. Colonies occur in 
grassy open meadows, in the alpine belt. Although the 
plants may be somewhat atypical of the California L. lyallii 
subsp. lyallii var. lyallii specimens, they are much like those 
of Oregon and Washington. 

1b. Lupinus lyallii subsp. lyallii var. danaus (A. Gray) S. 

Wats., Proc. Amer. Acad. 8: 534. 1873. 

L. danaus Gray, Proc. Amer. Acad. Arts & Sci. 7: 335. 
1868. Type: CALIFORNIA:  TUOLUMNE CO.: Mt Dana, 
Bolander 5087 (Holotype: GH; Isotype: UC, US). 

L. danaus var. bicolor Eastw., Leafl. West. Bot. 2: 201. 
1939. Holotype: CALIFORNIA: TUOLUMNE C0.: Mt Dana, 
Howell 14539 (CAS). Photos: (RSA, UMO). 

L. oreocharis Eastw., Leafl. West. Bot. 3: 171. 1942. 

430 Rhodora [Vol. 76 

Holotype: CALIFORNIA: TULARE CO.: Little Five Lakes 
Basin, Howell 17392 (CAS). Photos: (RSA, UMO). 

Plants differing from L. lyallit subsp. lyallii var. lyallii 
by matted vegetative growth, 1-4 cm. tall; plant overall 
height usually less than 10 cm. tall; pubescence frequently 
sparse, spreading; largest leaflets mostly 4-6 mm. long, 
rarely to 10 mm. long; peduncles 1.5-3 em. long, rarely to 
8 em. long; racemes capitate, 1.5-2.0 cm. long; flowers fre- 
quently more loosely arranged than in var. lyallii, smaller, 
6-7 mm. long. 

Representative specimens: California: FRESNO C0.: Humphreys 
Basin, headwaters Piute Creek, W of Mt Humphreys, Sharsmith 3136 
(uc). INYO co.: Piute Pass, E slope, Peirson 2552 (RSA). MADERA 
co.: Isberg Pk, 1 mi SW, Akey 332 (UC). MARIPOSA CO.: Mt Dana, 
Congdon, Aug 1898 (Uc). MODOC CO.: Mt Warren, Payne 307 
(JEPS). MONO C0.: San Juaquin Peak, Wright K3 (POM). TULARE 
co.: Mt Whitney, Clemens, Jul 1910 (POM, UMO). TUOLUMNE CO.: 
Gaylor Lakes, Howell 20401 (ws). 

The depauperate forms of L. lyallii subsp. lyallii are 
interpreted here as being ecological variants. Under a par- 
ticular set of environmental conditions populations of L. 
lyallii dwarfs to the matted form of var. danaus. However, 
frequently within the same population typical var. lyallii 
may occur in the larger form. Both varieties are found 
in high alpine rocky, sunny areas. Although this depau- 
perate or ecological variant of L. lyallii appears to be 
morphologically blending with the larger variety, it is 
considered here as a distinct taxon at the varietal level. 

A variant of similar vegetative morphology was observed 
among the Lupinus lyallii specimens of Canada and the 
higher alpine areas of Washington and Oregon, but the 
reproductive structures differ from that of var. danaus in 

Lupinus oreocharis is reduced to synonymy under var. 
danaus. Although some of these plants range to 14 cm. tall 
and resemble var. villosus in pubescence, and some may 
differ from var. danaus by having slightly elongated stems 

1974] Genus Lupinus — Cox 431 

which may have 1-2 cauline leaves, the available evidence 
does not indicate the warrant of a special epithet. 

1c. Lupinus lyallii subsp. lyallii var. fruticulosus (Greene) 

C. P. Sm., Bull. Torrey Bot. Club 51: 303. 1924. 

L. fruticulosus Greene, Muhlenbergia 8: 117. 1912. 
Type: OREGON: KLAMATH CO.: Anna Creek Valley, 
Austin 1650 (Holotype: US; Isotypes: NY, POM). Photos: 
(RSA, UMO, US). 

L. perditorum Greene, Muhlenbergia 8: 117. 1912. 
Holotype: OREGON: JACKSON CO.: Upper Rogue River, 
Applegate 2636 (US). Photo: (US). 

Plants in dense semi-shrubby clumps, branching pro- 
fusely, erect to semi-decumbent; stems 19-17 em: tall, 
usually less than 1.5 mm. in diameter; leaves cauline; 
petioles slender, 2-5 cm. long; leaflets 5-6, elliptic-oblanceo- 
late, largest 5-12 mm. long, av. 8.4, 1.8-3.5 mm. wide, length 
to width ratio 2.78-4.0, av. 3.42, subappressed, strigose- 
pilose on both surfaces; peduncles 2.5-5.5 em. long; racemes 
numerous, 1.5-4.3 em. long, 13-14 mm. in diameter, capitate 
and dense to somewhat scattered; flowers blue, 6.0-7.5 (8.0) 
mm. long, av. 6.9 mm.; bracts 1.5-3.0 mm. long; pedicels 
0.5-1.5 mm. long; banner elliptic to suborbicular, reflexed 
below the midpoint; pods 7.5-10 mm. long, 3.3-4 mm. wide; 
seeds 2-3. 

The greatest development of var. fruticulosus is along 
the flood plain of the Rogue River in Klamath Co., Oregon. 
Plants are found at elevations of 1067-1372 m., flowering 
in July. 

Representative specimens: Oregon: JACKSON CO.: Ashland, 9.7 km 
S, Sherwood 842 (WILLU). KLAMATH CO.: Crater Lake Natl. Park, 
Evinger 608 (osc). LINN CO.: Hunts Cove, Mt Jefferson area, 
Leach 4485 (ORE). 

Variety fruticulosus differs morphologically from other 
variants of Lupinus lyallii by its semi-shrubby base, pro- 
fusely branching clump habit and smaller flowers. A few 
L. lyallii specimens from Washington (mainly var. macro- 

432 Rhodora [Vol. 76 

florus) have thick, woody branching bases, but these are 
part of the underground caudex system and differ by having 
distinctively larger inflorescence structures. The Washing- 
ton material is more robust than fruticulosus, Variety 
fruticulosus is one of the most distinctive variations of 
L. lyallii subsp. lyallii. 

Variety fruticulosus is morphologically similar to var. 
roguensis, of the same general geographical region, and 
to subsp. alcis-temporis of the Clearwater and Selway 
Rivers in Idaho. It may be separated from these two taxa 
on the basis of its flower morphology, more scattered flower 
arrangement on the racemes and smaller size, as well as 
differences in vegetative traits. These plants exhibit a 
more erect woody branching habit with more filiform 
petioles and stems than any closely related entities. They 
are generally not as leafy, and the leaflets are fewer (5-6 
per leaf), smaller, and arranged in the upper portion of 
the branching clump. 

1d. Lupinus lyallii subsp. lyallii var. macroflorus B. J. Cox, 
var. nov. Type: WASHINGTON: PIERCE CO.: Mt Rainier, 
Yakima Park, Thompson 11086 (Holotype: MO; Isotypes: 
NY, POM-in part, US). Photo of holotype: (Cox 1972 a). 

Plantae perennes, humiles; caules ex caudice profuse 
ramificanti lignoso, 5-17 cm. alti, 1-15 mm. diametro, nodis 
0-2(8); petioli 3-5.5 em. longi; foliola 6-9, oblanceolata, 
maxima 5.5-11.3 mm. longa, 2.5-5.0 mm. lata; pedunculi 
3.0-5.5 em. longi, supra folia 1-2 em, extensi; racemi 2.5-5.5 
cm. longi, 2.5-3.0 cm. diametro, capitati usque subcapitati, 
floribus 5-17 in 1-5 verticillis; flores caerulei 10-12.5 mm. 
longi; calyx gibbosus; vexillum suborbiculatum usque obo- 
vatum ; ovula 3-5; legumina 13-15 mm. longa, 4-5.5 mm. lata. 

Plants perennial, in dense low mats; stems from a pro- 
fusely branching woody base, 5-17 cm. tall, 1-1.5 mm. in 
diameter, pubescence appressed to ascending, some spread- 
ing, cauline nodes 0-2(8); petioles 3-5.5 em. long; leaflets 
6-9, oblanceolate, largest 5.5-11.3 mm. long, av. 8.29 mm., 
2.5-5.0 mm, wide, av. 2.56 mm. ; peduncles 3.0-5.5 cm. long, 

1974] Genus Lupinus — Cox 433 

exceeding the foliage by 1-2 cm. or more; racemes 2.5-5.5 
em. long, 2.5-3.0 cm. in diameter, capitate to subcapitate, 
flowers 5-17 in 1-5 verticils; flowers blue, 10-12.5 mm. long, 
av. 11.2 mm. long; calyx gibbous at the base; banner sub- 
orbiculate to obovate, glabrous; keel densely ciliated near 
the acumen to glabrous in some; ovules 3-5; legumes 13-15 
mm. long, 4-5.5 mm. wide. 

Populations of var. macroflorus are found on the western 
side of the Cascade Mountain Range on both sides of the 
Columbia River in Washington and Oregon. They are found 
on Mary’s Peak in Benton Co., on Mt. Hood of Clackamus 
and Hood River Counties, Santiam Pass in Linn Co., Ore- 
gon, and Mt. St. Hellens of Skamania Co., and Mt. Rainier 
of Pierce Co., Washington. Like var. lyallii, var. macro- 
florus occurs at the upper vegetational limit between tim- 
berline and snowline at elevations of 1524-2378 m. Plant 
communities commonly associated include the species: 
Pinus contorta, P. albicaulis, Abies lasiocarpa, Tsuga. mer- 
tensiana and less frequently Pseudotsuga menziesii. 

Representative specimens: Oregon: BENTON CO.: Corvallis, W at 
Mary's Peak, Gilbert 108 (osc, US). CLACKAMUS CO.: Mt Hood, Jct 
of Hwys 26 & 35, Downey 4B (UMO, UTC). HOOD RIVER CO.: Mt Hood, 
Abrams 9445 (MO, POM, RM, WILLU). LINN CO.: Santiam Pass, 
Cronquist 7853 (GH, NY, RSA, UC, WS). Washington: PIERCE CO.: 
Mt Rainier Natl. Park, Burrough’s Mt & Frozen Lake, Flett 3121 
(WS). SKAMANIA CO.: Mt St Hellens, Buzzettii, Aug 1848 (osc). 

Variety macroflorus is distinct primarily on the basis of 
its larger flower size (10-12.5 mm. long) from var. lyallii 
flower size (7.5-10.5 mm. long). There are several inter- 
mediates between the variations of L. lyallii, but in general 
var. macroflorus is the extreme in flower morphological 

le. Lupinus lyallii subsp. lyallii var, roguensis B. J. Cox, 
var. nov. Type: OREGON: JACKSON co.: Farewell Bend 
Forest Camp, 1.6 km N of Union Cr, on Rogue River flood 
plain, Hitchcock & Martin 4997 (Holotype: RSA; Isotypes: 
MO, NY, UC, WS). Photo of isotype: (Cox 1972a). 

434 Rhodora [Vol. 76 

Plantae perennes, herbaceae, caespites densos ramificantes 
formantes, 2.5-3.5 dm. diametro, 12-18 cm. altae; caules 
humifusi usque erecti, 1-3 mm. diametro, ex caudice dense 
ramificanti lignoso, nodis 6-10, ad nodos ramificantes ; foliola 
7-9, maxima 8-13 mm. longa, 2.5-3.0 mm. lata; racemi capi- 
tati usque subcapitati 1.5-3.5 mm. longi, 4.5 mm. longi 
fructiferi, 1.9-2.5 mm. diametro; flores 7-9.5 mm. longi; 
vexillum et alae glabrae. 

Plants perennial, herbaceous, in dense branching clumps, 
2.5-3.5 dm. in diameter, 12-18 cm. tall; stems procumbent 
to erect, 1-3 mm. in diameter, from densely branching 
woody base, nodes 6-10, branching at the nodes; leaflets 
7-9, largest 8-13 mm. long, 2.5-3.0 mm. wide; racemes capi- 
tate to subeapitate, 1.5-3.5 mm. long, 4.5 mm. long in fruit, 
1.9-2.5 mm. in diameter; flowers 7-9.5 mm. long; banner 
and wings glabrous; keel ciliated along the upper margin 
near the acumen. 

Variety roguensis is found at Diamond Lake, Huckleberry 
Mt., Davis Lake, and on the flood plain of the Rogue River, 
in Jackson and Klamath Counties, Oregon, These plants 
occur in sandy soil in pine forests at elevations of 915-1067 
m. Flowering is from mid-June to mid-July. 

Representative specimens: Oregon: JACKSON CO.: Huckleberry 
Mt, Rogue River Natl. Forest, Thompson 130835 (MO, RSA, UC, WILLU, 
WS). KLAMATH CO.: Davis Lake, W shore, Hitchcock & Martin 4935 

Morphologically, these plants are most closely related to 
var. fructiculosus, but they differ in several significant 
characteristics. They are more robust, have larger branch- 
ing procumbent to erect stems, have larger flowers that 
are more densely arranged in racemes, and have more 
leafy foliage. The leaves are more numerous and are scat- 
tered throughout the clump, and the leaflets are larger 
with more leaflets per leaf. The manner of branching re- 
sembles that of L. aridus, especially subspecies lenorensis 
(Cox 1972a) found along the Clearwater and Selway Rivers 
in Idaho, Although the general branching pattern is often 

1974] Genus Lupinus — Cox 435 

difficult to detect, careful scrutiny finds verticillate spread- 
ing branching from an erect shoot; these branches become 
somewhat decumbent, but secondary and tertiary branches 
arise from them either adaxially or twisting around the 
decumbent stems and growing upward forming erect 

Several color variations are found in this segment of 
Lupinus lyallit. Many plants of var. roguensis have albino 
flowers (white to cream color — Maguire & Holmgren 26493 
— GH, NY, UC, US) ; however, others within the same popu- 
lation are intermediate in flower color to the albino and 
typical blue form (white to cream colored petals except for 
the purple banner and keel tip — Maguire & Holmgren 
26494 — GH, MO, NY, UC, US, WS). 

1f. Lupinus lyallii subsp. lyallii var. villosus Jeps., Jepson, 
Fl. Calif. 2: 267. 1936. Holotype: CALIFORNIA: ALPINE 
CO.: Carson Pass, Jepson 8100 (JEPS). 

L. hypoleucus Greene, Leafl. Bot. Obs. & Crit. 1: 74. 
1904. Type: CALIFORNIA: White Chief Peak, Culbert- 
son 16, Jul 1904 (distributed by Baker 4416 — specimen 
not located.) Paratype: CALIFORNIA: Mt, Goddard, 
Hall & Chandler 707 (US). 

Plants densely caespitose; stems decumbent to semi-erect, 
from a thick branching woody caudex, sericeous with as- 
cending to long spreading white hairs turning reddish- 
brown with age; petioles 4-5 cm. long; leaflets 6-7, elliptic- 
oblanceolate, apex acute-mucronate, largest 10-13 mm. long, 
3-5 mm. wide, villous on both surfaces; peduncles 3-7 cm. 
long; racemes densely verticilate 2.5-5 cm. long; bracts 
persistent, 4-6 mm. long; pedicels 1-1.5 mm. long; banner 
and wings glabrous; keel ciliate near the acumen; pods 
10-11 mm. long, 4 mm. wide. 

This variety occurs scattered throughout much of the 
distributional range of L. lyallii in several counties of 
California in the Sierra Nevada northward to several popu- 
lations in Klamath Co., Oregon. 

436 Rhodora [Vol. 76 

Representative specimens: California: ALPINE CO.: Carson Pass, 
Higgins 1752 (BRY). FRESNO CO.: Bubbs Creek, Peirson 531 (RSA). 
HUMBOLDT CO.: Grouse Mt, back of Clear Lake, Tracy 16672 (UC). 
LASSEN C0.: Grass Valley, Fisher & Johnson F203 (UC). MADERA 
co.: Garnet Lake, Howell 16421 (RSA, UC, US). MARIPOSA CO.: 
Donohue Pass, Schreiber 1767 (UC). MENDOCINO CO.: South Yolla 
Bolly, Jepson 57t (JEPS). MONO CO.: San Juaquin Mt, Howell 
16875 (Ws). PLACER CO.: Mt Lincoln, Howell 18670 (US). SHASTA 
Yolla Bolly, Jepson 57t (JEPS). MONO C0.: San Juaquin Mt, Howell 
16875 (ws). PLACIER CO.: Mt Lincoln, Howell 18670 (US). SHASTA 
co.: Squaw Valley Creek Meadows, Cooke 13651 (NY). TRINITY CO.: 
Trinity Center, 3 mi S, E of Scott Ranch, Cantelow, Aug 1936 (RSA). 
TUOLUMNE C0.: Gaylor Lakes, Howell 20418 (GH). Oregon: KLA- 
MATH CO.: Llao Rock, Baker 6245 (NY, OSC, UC, WS). 

Variety villosus is recognized primarily on the basis of 
its dense pubescence. In L. lyallii, pubescence ranges from 
puberulent-sericeous to woolly-villous approaching spread- 
ing hirsute. Several populations exhibit different forms of 
pubescence within a single colony; this is especially true 
of those specimens viewed from the Carson Pass vicinity 
(the type locality of var. villosus), Alpine Co., California. 

2. Lupinus lyallii subsp. alcis-temporis (C. P. Sm.) B. J. 

Cox, comb. nov. 

Lupinus alcis-temporis C. P. Sm., Sp. Lup. 558. 1946. 
Type: IDAHO: CLEARWATER CO.: Pierce, 17.7 km. E, 
Crystal Creek Grade, Christ 14634 (Holotype: DS; Iso- 
types: NY, US). Photos: (RSA, UMO). 

Plants sprawling caespitose, branching profusely from 
a slender woody base, in clumps 11-30 cm. in diameter; 
stems subdecumbent, 8-15 em. long, strigose-sericeous with 
dense coarse ascending hairs, nodes 4-6; cauline petioles 
2-4 em. long; stipules 7-9 mm. long, connate to the petioles 
3-4.5 mm., filiform; leaflets 5-7, elliptic-oblanceolate, apex 
acute to short mucronate, longest leaflets 2-18 mm., av. 
10 mm. long, 0.8-5 mm. wide, length to width ratio 2.33- 
4.33, av. 3.18, strigose-pilose to villous on both surfaces, 
some more sparsely so on the upper surface to glabrous; 
peduncles 1.5-3.5 cm. long; racemes capitate, 2-3.5(6) cm. 
long, densely verticillate; flowers blue with yellow sulcus, 

1974] Genus Lupinus — Cox 437 

7-8.2 mm. long; bracts 5-9 mm. long, subulate-attenuate, 
sericeous to villous; pedicels 1.2-1.8 mm. long; calyx upper- 
lip 4.0-5.0 mm. long, bilobed, notch 2-3.5 mm. deep, lower- 
lip 5.0-6.0 mm. long, apex tridentate, center tooth 0.2-0.5 
mm. long, lips connate 1.0-1.5 mm. ; bracteoles 0.8-1.5 mm. 
long, attached at the sinus, villous ; banner elliptic-oblanceo- 
late, 7.0-8.0 mm. long, 4-4.5 mm. wide, length to width 1.56- 
1.88, av. 1.74, reflexed to appressed ratio 1.25-1.50, av. 1.37, 
angle 148-158°; wings 6.0-7.3 mm. long, 3.0-3.4 mm. wide, 
length to width ratio 1.97-2.85, av. 2.16; keel 2 mm. wide 
at the widest point, angle 111-122°, ciliated sparsely near 
the acumen ; ovules 2-3; pods 8.5-9.5 mm. long, 3.4-4.0 mm. 
wide, densely short woolly; seeds 2-3, 1.8-2.0 mm. X 2.0- 
2.2 mm., 0.9-1.1 mm. thick, tan color. 

Subspecies alcis-temporis is found in sandy and gravelly 
slopes on road cuts, generally in otherwise barren areas. 
One population occurs in a Pinus monticola forest; the 
Christ & Smith collection (15385 — NY) was found on 
“forest soil bordering beds of Boykinia major; the Dauben- 
mire collection (47150 — WS) was made in a Spruce-Fir 
Zone. Plants oecur at 915-1220 m. elevation. 

Representative specimens: Idaho: CLEARWATER CO.: Pierce, 17.7 
km E at the confluence of Big Breakfast Creek and Crystal Creek, 
Christ & Smith 15385 (NY); Washington and Lodge Creeks, S31, 
T39N, R7E, Lingenfelter 466 (NY, RSA, UC, WS). 

This taxon was first described by C. P. Smith from a 
Christ “little number picked up in Clearwater County 
(Idaho) during elk season in October.” Hence the name 
*Christ's Elk-Season Lupine”, Lupinus alcis-temporis in 
Smith’s Latinum utilizes the genitive for both elements of 
the compound name (alces — elk and tempus — season). 
Both Christ and Smith were deeply interested in the fact 
that a lupine in anthesis could be found in Clearwater 
Co. in the middle of October “without a trace of fruit or 
seed, and at only 4,000 feet elevation." 

Several stages of development are found in more ex- 
tensive study of other collections of these lupines, all of 

438 Rhodora [Vol. 76 

which have been collected on July 10th or July 11th. Small 
clusters of young herbaceous-stemmed, procumbent plants 
may be found flowering the same time as older plants with 
semi-woody branching stems, sprawling 30-45 cm. in di- 
ameter. The isotype material viewed from NY and US 
appears merely to be atypically late in reaching anthesis. 
It is in anthesis with some fruit in the initial stages of 
enlargement, and all branches are woody including second- 
ary and tertiary branches; only the leaves and inflorescences 
remain herbaceous at this late seasonal date. 

3. Lupinus lyallii subsp. minutifolius (Eastw.) B. J. Cox, 
comb. & stat. nov. 
Lupinus minutifolius Eastw. Leafl. West. Bot. 2: 267. 
1940. Holotype: OREGON: HARNEY CO0.: Fish Lake, 
Steens Mts., Henderson 8132 (CAS). Photos: (RSA, UMO). 

Plants acaulescent; foliage to 4.5 cm. tall; stems 4-6 cm. 
tall, appressed hispidulose with more coarse scaberose 
hairs; petioles filiform, 2.5-4 cm. long; leaflets 5-7, elliptical, 
glabrous above to sparse pubescence on the margins, short 
coarse appressed hairs below, largest 6-9 mm., av. 6.8 mm., 
2.9.8 mm. wide, av. 2.45 mm., length to width ratio 2.4-3.0, 
av. 2.75; peduncles 6-8 cm. long or only 3 cm. long in im- 
mature plants; racemes 1.5-3.0 em. long, 3-4 compact verti- 
cils, ascending or erect from nearly prostrate stems; flowers 
7.5-10 mm. long, mostly blue with nearly white banners 
and purple tipped keels; bracts caducous, 4.5-6 mm. long, 
subulate; pedicels 1.0-2.0 mm. long; calyx upper lip 4-5 mm. 
long, bilobed, notched 2.5-2.7 mm. deep, lower-lip 4.5-5.0 
mm. long, apex entire-serrate to tridentate with the center 
tooth approaching 0.7 mm. long, lips connate 1.2-1.3 mm., 
bracteoles 0.2 mm. long, attached at the sinus, strigose to 
short ascending pilose; banner elliptic-oblong, 8-8.5 mm. 
long, 4.2-5.5 mm. wide, length to width ratio 1.45-1.90, av. 
1.65, reflexed to appressed ratio 1.08-1.57, av. 1.34, angle 
150-160°; wings 8.8-9.0 mm. long, 3.8-4.2 mm. wide, length 
to width ratio 2.10-2.57, av. 2.34, claws 1.0-1.8 mm. long, 
glabrous; keel 2.0-2.2 mm, wide at the widest point, angle 

1974] Genus Lupinus — Cox 439 

118-1235, ciliated along the outer half of the upper margin, 
densest toward the acumen; ovules 3; pods 9.0-10.5 mm. 
long, 3.5-4.2 mm. wide, short appressed; seeds 2-4. 

Subspecies minutifolius is found in the Steens Mountains 
of Harney Co. Oregon, geographically replacing subsp. 
lyallii. These plants are found on rocky, gravelly, gentle 
basaltic slopes but frequently occur in alpine meadows on 
hillside areas with grasses and sedges in sloping rocky 
habitats. Several specimens have been collected in beds of 
lingering snowbanks where snow has accumulated and 
remained during the winter and spring. Anthesis generally 
occurs during the latter part of July. Plants are found at 
elevations of 2439-2896 m. 

Representative specimens: Oregon: HARNEY CO.: Big Fish Creek 
Canyon, Maguire & Holmgren 26789 (GH, NY, UC, US); Frenchglen, 
31.4 km due E-SE, Hansen 490 (osc). 

Although some populations of subsp. minutifolius re- 
semble subsp. lyallii var. danaus, they have several distinct 
morphological characteristics. The flowers of subsp. minu- 
tifolius generally have more oblong shaped banners, more 
linear wings, and therefore a greater length to width petal 
ratio than subsp. lyallii. Although the vegetative mor- 
phology of subsp. minutifolius resembles the alpine var. 
danaus and other northern forms of subsp. lyallii, it shows 
distinction in the filiform petioles from the basal clumps 
of less than 4.5 cm. tall, and frequently by the fewer (4-7, 
av. 5) leaflets that are glabrous on the upper surface. The 
plants of the Cronquist collection. (8694 — NY, OSC, RM, 
RSA, UC, WS) are morphologically larger than normal for 
the taxon. The foliage of these specimens approach 5 cm. 
tall, overall plant height approaches 9.5 cm.; flowers are 
an intermediate 8.5 mm. long ; the leaflets are larger 10 mm. 
x 8.5 mm.; peduncles are larger 6.5-7 mm. long, and 
racemes approach 4 em. long. 

4. Lupinus lyallii subsp. subpandens C. P. Sm. ex Dunn, 
El Aliso 3: 171. 1955. Type: NEVADA: ELKO CO.: Elko, 

440 Rhodora [Vol. 76 

113 km. NW, Holmgren, Jun 1938 (Holotype: UTC; Iso- 
types: NA, NEV, AGR.). Photos: (RSA, UMO). 

Plants with many small tufts from a profusely branch- 
ing woody caudex, erect; foliage 4.5-9.0 cm. in height, 
mostly less than 6 cm. tall; stems 7-18 cm. tall, usually less 
than 1 mm. in diameter, appressed strigose-pilose pubes- 
cence less than 2 mm. long ; leaves in basal clusters ; petioles 
slender, 3-9 cm. long; stipules inconspicuous, 6-10 mm. long, 
connate to the petioles 4-5 mm, subulate; leaflets 5-6, linear- 
oblanceolate, apex acute, largest 9-22 mm. long, 2-5 mm. 
wide, length to width ratio 3.5-6.3, av. 4.8, sericeous-strigose 
below, more sparsely so above with glabrous apex and mid- 
rib; peduncle 5-9 cm. long, usually exceeding the foliage by 
2-5 cm.; racemes capitate to subcapitate, 1.5-4 cm. long; 
flowers dense, blue, 9.5-11.5 mm. long; bracts 4-8.5 mm. 
long, appressed to ascending pilose; pedicels 1-2 mm. long; 
calyx upper-lip 4.2-5.5 mm, long, notch 1.3-3.2 mm. deep, 
lower-lip 4.5-5.5 mm. long, apex deeply serrate to tridentate, 
center tooth to 0.5 mm. long, lips connate 1.0-1.5 mm., 
bracteoles slender, inconspicuous, 0.5-1.0 mm, long, attached 
to the sinus, strigose-sericeous, hairs to 2 mm. long; 
banner elliptic, 8-9.5 mm. long, 5-6.5 mm. wide, length to 
width ratio 1.45-1.90, av. 1.56, reflexed to appressed ratio 
1.13-1.71, av. 1.44, angle 146-150°; wings 8.2-9.8 mm. long, 
3.4-4.8 mm. wide, length to width ratio 2.04-2.65, av. 2.36, 
claws 1.5-2.0 mm. long, glabrous; keel 2.2-3.0 mm. wide at 
the widest point, angle 108-120°, ciliated along the outer 
half of the upper margin, most densely near the acumen, 
cilia to 0.7 mm. long; ovules 3-5; pods 11-14 mm. long, 
5-5.5 mm. wide, dense strigose to short villous; seeds 2-3, 
2.0-2.2 mm. X 2.8-3.0 mm. 1.3 + 0.1 mm. in thickness, 
beige to tan with darker brown mottling. 

These plants are found in dry, rocky creek beds and 
open well-drained draws in northeastern Nevada, in Elko 
Co., and southwestern Idaho, in Camas and Owyhee Coun- 
ties. The most common associate is Artemisia sp. Plants 
flower from mid-July to the first part of August. Altitudi- 

1974] Genus Lupinus — Cox 441 

nally, they are found at 1677-1982 m., with one population 
found on Alleghany Creek at 2439 m. 

Representative specimens: Idaho: CAMAS CO.: Fairfield, 16 km. 
SE, Ripley & Barneby 10671 (NY). OWYHEE Co.: Mud Flat, 4.8 km. 
SW, on road to Juniper Mt, Maguire & Holmgren 26305 (GH, POM, 
UC, US, UTC, WS). Nevada: ELKO Co.: O’Neil, 16 km. S, Holmgren, 
Cooper & Cain 181 (GH, osc, UTC) ; Wells, 48 km. N, Jones, Jul 1901 

Dunn (1955) interpreted Lupinus lyallii subsp. sub- 
pandens as an acquisition of a combination of the char- 
acters of L. aridus with those of L. lyallii. The pubescence 
type and system of branching from the caudex, as well 
as certain flower morphology, is characteristic of L. aridus, 
whereas the small caespitose habit with elongated peduncles 
and capitate inflorescences with small flowers is more in- 
dicative of L. lyallii. Dunn further suggests that the plant’s 
occupancy of the flatland Artemisia habitat of 1677-1982 m. 
elevation indicates a considerable amount of the L. aridus 
physiological adaptations, contrasting the higher altitudinal 
L. lyallii habitat. 

This study concurs with Dunn's interpretation of the 
affinity of subsp. subpandens with L. lyallii. These plants 
inhabit a geographically distinct region, also having both 
morphological and physiological distinction. Although each 
subsp. subpandens population has aequired a unique com- 
bination of characteristics, the plants can usually be recog- 
nized by the erect habit of both flowering culms and petioles 
as well as by the glabrous apex and midrib of the upper 
leaflet surfaces. 

The specimens from Camas Co., Idaho (Ripley & Barneby 
10671 — NY) have larger flowers than the taxon mean, with 
banners only slightly reflexed and frequently with one node 
on the flowering culm above 2 cm. from the basal cluster 
of leaves, and racemes of 3-5 cm. long, being about 2 cm. 
in diameter. 

The specimens collected south of O'Neil, Nevada (Holm- 
gren, Cooper & Cain 181 — GH, OSC, UTC) are more slight 
in habit, having shorter petioles with smaller leaflets that 

442 Rhodora [Vol. 76 

are more glabrous on the upper surface, and shorter, more 
capitate racemes with the banners less reflexed. 

The Mud Flat, Owyhee Co., Idaho specimens (Maguire 
& Holmgren 26305 — GH, POM, UC, US, UTC, WS) have fewer 
leaflets than other populations, and the petioles are more 
elongated and slender. 

5. Lupinus lyallii subsp, washoensis (Heller) B. J. Cox, 

comb & stat. nov. 

L. washoensis Heller, Muhlenbergia 6: 72. 1910. based 
on L. pinetorum Heller, Muhlenbergia 6: 25. 1910, not L. 
pinetorum Jones, Contr. West. Bot. 8: 25. 1898. Type: 
NEVADA: WASHOE CO.: Ridge S of Alum Creek, Heller 
9748 (Holotype: Nev. Agr.; Isotypes: DS, GH, ISC, MO, 
NY, ORE, POM, UC, US, WS). Photos: (RSA, UMO). 

L. aridus var. washoensis (Heller) C. P. Sm,, Bull. 
Torrey Bot. Club 51: 3083. 1924. 

Plants 10-18 cm. tall, in dense caespitose clumps, 15-30 
cm. in diameter, from woody vertical caudex branching at 
ground level, with 10-18 racemes flowering simultaneously; 
stems procumbent to erect, 1-2.5 mm. in diameter, villous 
to hirsute-shaggy white pubescence 2-4 mm. long; 2-3 
nodes; cauline petioles 4.5-8 cm. long, pubescence as on the 
stems; basal petioles on immature plants 4.5-7 cm. long; 
stipules 6-10 mm. long, connate to petioles 4-6 mm., densely 
white bearded pubescence; leaflets 5-6, oblanceolate to ellip- 
tic-oblong, apex short mucronate, largest 9-21 mm. long, 
av. 13.7 mm., 3-7 mm. wide, length to width ration 1.86- 
3.27, av. 2.71, silvery appressed to ascending pubescence 
below, more sparsely so above, upper surface appearing 
more greenish; peduncles 3-5 cm. long, generally exceeding 
the leaves; racemes 3-6 cm. long, 1.6-1.9 cm. in diameter, 
densely subverticillate; flowers blue to violet, banner with 
a white sulcus, 8-9 mm. long; bracts 3-5.5 mm. long, subu- 
late-attenuate, pilose-sericeous; pedicels 1.5-2.0 mm, long; 
calyx upper-lip 3.5-4.5 mm. long, bilobed, the notch 0.8-1.2 
mm. deep, lower-lip 4-5.5 mm, long, apex serrate, lips con- 
nate 1-2 mm., bracteoles 0.5-1.0 mm. long, attached at the 

1974] Genus Lupinus — Cox 443 

sinus, villous; banner elliptic-oblong, 7.8-8.0 mm. long, 5.0- 
6.0 mm. wide, length to width ratio 1.33-1.67, av. 1.52, 
reflexed to appressed ratio 1.25-1.35, angle 140-150? ; wings 
7.5-8.0 mm. long, 4-4.5 mm. wide, length to width ratio 
1.74-2.00, av. 1.87, claws 1.5-2.0, glabrous; keel 2.5-3.0 mm. 
wide at the widest point, angle 118-125°, ciliated along the 
upper margin near the acumen; ovules 2-3; pods 9-13 mm. 
long, 5-6 mm. wide, villous to ascending woolly; seeds 1-3, 
1.8-2.2 mm. X 2.4-2.7 mm., 0.9-1.2 mm. thick, tan with 
brown mottling. 

Subspecies washoensis has a restricted habitat on the 
eastern slopes of the Sierra Nevada in the southern part 
of Washoe Co., SW of Reno, Nevada. Plants occur on 
gravelly montane open pine slopes at elevations of 1524- 
2134 m., flowering in June. 

Representative specimens: Nevada: WASHOE CO.: Dinsmore Camp, 
Hunter Creek Canyon, Kennedy 1614 (Mo, Ny, UC); Reno, Hillman 
(POM); 10-13 km. SW, Hunter’s Creek Road, Archer 6090 (NY). 

The general appearance of some of these plants indicates 
relationship with Lupinus aridus, a portion of the racemes 
being exceeded in height by the foliage. Affinity is also 
indicated with L. sellulus var. lobbii in general morpho- 
logical structure (Cox 1972a, 1973c). Considering the 
total conformation — narrow, short subcapitate or longer 
racemes, and the procumbent, branching, sprawling vege- 
tative habit — this study concludes that the washoensis 
affinity lies most closely with L. lyallii. These plants re- 
place L. lyallii subsp. lyallii geographically and ecologically, 
aso being recognizably distinct by somewhat larger stature, 
having longer racemes, larger leaves, more yellow-green 
color and spreading silvery stem and petiole pubescence. 

The Dinsmore population (Kennedy 1614 — MO, NY, UC), 
which is only a few miles north of the type locality in 
Hunter Creek Canyon, has been used in sheep feeding ex- 
periment (Nev. Agr. Exp. Sta. Bull. 71: 30. 1909). The 
plants are said to be relished by the sheep, proving to be a 
good forage plant. 

Rhodora [Vol. 76 

` oo N 
———— \ 
o% E 
a |! 
is we RN i ( 
* & i 
/ 1 
, \ 
/ IDAHO | 
^ | 
q—. de. | 
: Ap ——— 
| AA! 
b | subspecies: variety: 
A | 
odes. | o-lyallii ——— lyallii 
33 N j +- lyallii ————— fruticulosus 
3 x 
EAT X | *- | yallii villosus 
Et i i e - | yallii roguensis 
oo N M 
go NOM x—lyallii danaus 
ii .. CALIFORNIA N v - lyallii macroflorus 

A - washoensis 
a- minutifolius 
x - alcis- temporis 


Figure 1. Distribution of Lupinus lyallii. 

1974] Genus Lupinus — Cox 445 


Cox, B. J. 1972a. Biosystematies of Lupinus lepidus-L. caespitosus 
complex. Ph.D. dissertation. Univ. of Missouri, Columbia. 343 p. 
1972b. IOPB Chromosome number reports XXXVIII. 
Taxon 21: 680-681. 
1973a. A chemosystematic comparison of the Lupinus 
lepidus-L. caespitosus complex. Bull. Torrey Bot. Club 100: 12-17. 
1973b. Protein relationships among the perennial cae- 
spitose lupines. Bull. Torrey Bot. Club (in press). 
1973c. Taxonomy of Lupinus sellulus (Papilionaceae). 
Amer. Mid. Natur. (in press). 
Dunn, D. B. 1955. A new subspecies of Lupinus lyallii. El Aliso 

$2 2171-172. 
PHILLIPS, L. L. 1957. Chromosome numbers in Lupinus. Madroño 
14: 30-36. 




While collecting vascular aquatie plants in several ponds 
in east-central Massachusetts we found that there was a 
considerable accumulation of algae upon some of the vascu- 
lar species, but very little on others. The present paper is 
the result of our futher observations on and examination of 
the relationship between the algae and the vascular plants 
upon whose surfaces they were found. 

There is little published data elaborating the substrate 
preferences of fresh-water algae. Water chemistry, light, 
temperature, water density, pH, and a wide variety of 
metabolic processes have been treated frequently (see Jack- 
son, 1964, and Smith, 1950) as environmental factors in 
algal growth. Reference texts (e.g., Prescott, 1962) indi- 
cate that some species of algae are epilithic, endophytic, or 
epizooic, or that many are epiphytic upon vascular plants or 
larger algae. Other than Chapman (1962) there seems to 
be no treatment of situations in which the algae have the 
opportunity to grow on or in association with a variety of 
substrates and in which clearly some means of preference 
for one another is established. Tiffany (1951), for exam- 
ple, suggested that the amount of substrate available to the 
spores for fixation is of major importance to algal growth, 
but he fails to include any discussion of the character of 
the substrate itself, or of the various environmental factors 
that might be operative in the selection and utilization of a 
particular substrate. Chapman suggests that, (1) the age 
and nature of the substrate flora are important, (2) diatoms 
are more likely to be on older leaves, (3) epiphytes are most 
numerous on well lighted surfaces, (4) epiphytes are most 
numerous on submerged material, (5) depressions between 
cells of the host surface seem to provide better substrate for 
attachment, and (6) ponds with muddy bottoms have a 


1974] Algae on Haloragaceae — Colt & Hellquist 447 

reduced number of epiphytes. With few epiphytes occur- 
ring in our collections we tend to agree with the last prem- 
ise, but we have insufficient evidence to either support or 
refute the other ideas. We might generally suggest, how- 
ever, that rather similar principles may operate for both 
tychoplankters and epiphytes. 

Our examination of material from different ponds clearly 
showed that algae grew profusely upon some vascular plants 
but sparsely or rarely on others. We also found that both 
the number of species and the types of tychoplankter algae 
varied with similar plants in different environments. In 
order to identify, if possible, the operative environmental 
factors, we selected two sites in which the algal habitats 
might be contrasted. In both instances, (1) a small leaved 
aquatic plant was the primary algal substrate, (2) the sub- 
strate plant grew in the shallower portions of the pond, 
(3) both ponds and collecting sites were in virtually iden- 
tical locations beside highways, and (4) there was no shad- 
ing of the collecting stations by macrovegetation on or near 
the banks of the pond. 


1. Pratt Pond. This pond is located at the junction of 
Grove Street, Hopkinton Road, and Westboro Road in the 
town of Upton, Worcester County, Massachusetts. 

All collections were made in the western sector of Pratt, 
from water varying between 1.5 and 2.0 meters in depth. 
Most of the remainder of the pond is sufficiently deep so as 
to permit swimming and diving. The bottom of the pond is 
sandy-mud throughout. The pH averaged 6.5 and the alka- 
linity was 3 ppm. 

The dominant vascular aquatic in the collection area of 
Pratt was Myriophyllum heterophyllum Michx., occurring 
over more than one-third of the center of the bay. In so far 
as we were able to determine during the several months of 
field work, no other vascular plant grew in or near the stand 
of Myriophyllum. This resulted in a clear zone around the 
stand extending to within several meters of the pond’s edge. 

448 Rhodora [Vol. 76 

In this shallower water a few other rooted aquatics grew 
sparsely. Of these Nymphaea odorata Ait., Pontederia cor- 
data L., and Potamogeton spirillus Tuckerm., were the most 

2. Icehouse Pond. This pond is beside West Main Street, 
0.9 kilometers east of Interstate Route 495, in the town of 
Hopkinton, Middlesex County, Massachusetts. 

Samples were collected from water up to 0.5 meters in 
depth along the southern edge of the pond. The deepest 
portion of this pond was approximately 1.5 meters, the 
bottom being mud and gravel. Water loss from this pond 
during the late summer months left some of the westerly 
edges as exposed mud and rocks. The pH averaged 6.1, and 
the alkalinity was 3.5 ppm. 

The shallow waters of this pond were densely populated 
with a variety of rooted aquatic plants. Algae were asso- 
ciated most profusely with Proserpinaca palustris L., which 
grew in scattered stands of a few plants each. Other com- 
mon vascular plants in the collection area were Potamogeton 
pulcher Tuckerm., P. capillaceus Poir., Eleocharis acicularis 
(L.) R. & S., Pontederia cordata f. taenia Fassett, Nym- 
phaea odorata Ait., Nuphar variegatum Engelm., Brasenia 
sehreberi Gmel., Ludwigia palustris var. americana (DC.) 
Fern. & Grisc., Callitriche verna L., Myriophyllum humile 
(Raf.) Morong, and Utricularia purpurea Walt. 


samples of the vascular plants were collected from the 
two pond stations and the species determination made using 
Fassett (1966) and Fernald (1950). Representative speci- 
mens were pressed and are in the herbarium at Boston 
State College. 

samples of the primary substrate plants were collected, 
some portions retained fresh for immediate scrutiny, and 
the remainder preserved in a solution of 3% formalin. De- 
termination of the associated algal species was made from 
Smith (1924), Irenee-Marie (1939), Prescott (1964), and 
Patrick & Reimer (1966). Microscope slides were made of 

1974] Algae on Haloragaceae — Colt & Hellquist 449 

some algae using the Kaiser Glycerin-Gelatin method in 
order to facilitate drawing by camera lucida. All other 
samples were placed in Transeau's Fluid for permanent 


Twenty-one species of algae were found on or in asso- 
ciation with Myriophyllum heterophyllum in Pratt Pond. 
Sixty-one algal taxa were discovered in various degrees of 
association with Proserpinaca palustris in Icehouse Pond. 
Seven species were common to each pond. The wide dis- 
parity in numbers despite virtually simultaneous collection 
times strongly suggests that Proserpinaca offers a more 
favorable habitat to the algae than does Myriophyllum. In- 
asmuch as a difference in surface characteristies might 
lead to the difference in habitats, it was the first possibility 
examined. We were not able to substantiate such a premise 
on any physical basis. One of the problems, however, is that 
only five genera of epiphytes were recorded during the 
study. Stigeoclonium and Characium each had one species 
present infrequently in Pratt Pond, while Bulbochaete and 
Gloeotrichia each had one commonly found species in Ice- 
house Pond, and Oedogonium with one common species and 
one infrequently encountered, both in Icehouse Pond. The 
pattern of occurrence of these was generally the same as 
for the tychoplankters and appeared to be due to the overall 
differences in the habitats rather than individual differences 
between the substrate plants. 

Thus it seemed likely that there was a combination of 
factors operating in each situation, with sufficient variance 
between the two habitats such as to account for the differ- 
ence in algal populations. This view is strengthened by the 
difference in tychoplankter populations, Icehouse Pond hav- 
ing approximately twice as many taxa. Inasmuch as the 
tychoplankters do not adhere to a particular substrate, but 
live in association with the substrate, they would not neces- 
sarily be affected by physical characteristics. 

Some previous work gave direction to our scrutiny. An- 

450 Rhodora [Vol. 76 

derson & Walker (1920) concluded from an investigation 
of some shallow Nebraska lakes that light and mechanical 
support are of primary influence to algal growth. They also 
suggested that shallow water and wind protection give rise 
to as nearly uniform conditions as possible. Needham & 
Lloyd (1916) showed that in shallow waters with abundant 
vegetation currents are reduced or eliminated and there is 
little possibility of wind disturbance. The authors' own 
experience has been that tychoplankter populations are par- 
ticularly rich among the wet sphagnum of New England 

Comparison of the two sites on these bases yielded some 
rather interesting results. It has already been stated that 
sunlight was readily and equally available to the water in 
which both sets of substrate plants grew. The collecting 
area in Icehouse Pond, however, was much shallower than 
that of Pratt Pond. This difference was negated by the 
position of the tychoplankter algae in Pratt Pond; they 
were all within 14 meter of the surface of the water. The 
only living algae below that level in Pratt Pond were dia- 
toms, and they were usually obscured by the detritus on the 
lower branches of plants of Myriophyllum. 

Physically, Icehouse Pond is protected from surface winds 
by the mixed deciduous-coniferous forest around it. Al- 
though its total surface area approximates that of the sec- 
tion of Pratt Pond from which we collected, we were unable 
to observe more than mere surface ripples on Icehouse Pond 
during any visit to the site. Part of the reason for this is 
the dense vegetation within the water itself. At Pratt Pond, 
however, we were able to note considerable surface disturb- 
ance by even relatively light wind currents. 

We were unable to observe any current within Icehouse 
Pond. This is largely due to the dense aquatic vegetation. 
Water movement could easily be discerned in Pratt Pond. 
As we collected samples in Pratt Pond, broken pieces of 
Myriophyllum would rise within the stand of plants then 
drift “downwind.” Our observation of this phenomenon 
showed that the drag of the wind on the surface of the 

1974] Algae on Haloragaceae — Colt & Hellquist 451 

water caused the upper portion of the water to move, cre- 
ating an effective pull on the lower water body. Water then 
rose through the plants and downwind. Davis (1955) sug- 
gested that tychoplankters would be unable to adapt evolu- 
tionarily to such current conditions, On the basis of our 
limited work we concur, finding no evidence of specialized 
attachment structures or polymorphism which would give 
any species an advantage in maintaining its position. It 
appears to us that the lack of water movement within Ice- 
house Pond is conducive to an optimum interchange of 
chemical nutrients. On the other hand, the flow within 
Pratt Pond apparently reduces a similar interchange, thus 
inhibiting population growth to some degree. 

Our final consideration was the growth habit of the small 
leaved algal substrates and the associated macroflora. In 
Pratt Pond Myriophyllum produced vertical axes greater 
than the depth of the water. Near the surface of the water 
the distal portions of the plants would bend with the drag 
of the surface water. This appeared to produce a matting 
effect at the surface which would effectively reduce the light 
available to lower leveis. The Proserpinaca in Icehouse 
Pond grew only to the surface of the water and distal flex- 
ing was rare and then of a minor nature, Light could pass 
easily through the finely dissected leaves to lower levels. 
Moreover, the densely crowded vegetation of Icehouse Pond 
effectively reduced lateral “waving” by the plant. Pro- 
serpinaca occurred in larger clumps than any of the other 
small leaved plants in Icehouse Pond (e.g., Myriophyllum, 

Examination of the broader-leaved aquatics in each pond 
did not reveal any substantial amount of algae growing on 
or associated with them. The most frequent species were 
diatoms, Tabellaria, for example, being the most common. 


From our observations we conclude that for the optimum 
growth of tychoplankter algae there are at least three basic 
requirements. Not necessarily in order of importance they 

452 Rhodora [Vol. 76 

are: (1) a substrate which provides both maximum surface 
area and exposure to the available light; (2) an absence of 
strong or prolonged water currents; and (3) the presence 
of a varied flora which provides a broad spectrum of nutri- 
ents within the system. 

Based on our observations at both ponds we suggest that 
small leaved plants will most often provide the optimum 
habitat for tychoplankter algae. We are not convinced, 
however, that monospecific stands provide the optimum en- 
vironment for tychoplankters, although such stands may be 
the best of the available substrates. We have found com- 
parable situations in bogs, as it has been our experience 
that the sphagnum bogs with the greatest variety of macro- 
flora usually yield the largest number of algal species. 

We further suggest that pH and water temperatures are 
primarily useful only in contrasting regions with geograph- 
ical and/or geological differences. It can be reasonably 
inferred that evolution has already adapted local algal 
species to pH and water temperature as they occur in the 
algal habitat. 

Finally, we concur with previous workers who have sug- 
gested that quiet, stable and uniform water conditions are 
best for the growth of tychoplankter algal populations. 

The list of algae below follows the sequence of Smith 
(1950). Nomenclature of the Bacillariophyceae is according 
to Patrick & Reimer (1966). Location and frequency are 
indicated by the following: IP — Icehouse Pond, PP — 
Pratt Pond, R — rare (less than 5 collected in all samples), 
F — frequent (averaged up to 10 in all samples), C — com- 
mon (averaged more than 10 in all samples), E — epiphyte, 
T — tychoplankter. 

Stigeoclonium subsecundum Kuetzing PP-R-E 
Bulbochaete Furberae Collins IP-C-E 
Oedogonium michiganense Tiffany IP-C-E 
O. pseudoplenum Tiffany IP-C-E 

Characium ornithocephalum A. Braun PP-R-E 

1974] Algae on Haloragaceae — Colt & Hellquist 453 

Pediastrum Boryanwm (Turp.) Meneghini IP-R-T 

P. duplex var. cohaerens Bohlin 

P. tetras (Ehrenberg) Ralfs 

P. tetras var. tetraodon (Corda) 

Nephrocytium lunatum W. West 

Kirchneriella contorta (Schmidle) 
G. M. Smith 

Scenedesmus abundans var. brevicauda 

G. M. Smith 
S. bijuga (Turp.) Lagerheim 
S. dimorphus (Turp.) Lagerheim 

S. quadricauda var. maxima West & West 
- Actinastrum Hantzschii Lagerheim 
two easily separable species 

Spirogyra spp. 





were common in the vegeta- 
tive state in Icehouse Pond. 

Gonatozygon pilosum Wolle 
Triploceras gracile Bailey 
Closterium Jenneri Ralfs 
Cl. parvulum Naegeli 

Cl. Ralfsii var. hybridum Rabenhorst 

Cl. setaceum Ehrenberg 

Pleurotaenium coronatum (Breb.) 

P. subcoronulatum var. detum 
West & West 

P. Trabecula (Ehrenb.) Naegeli 

P. Trabecula var. rectum (Delp.) 
W. West 

P. Trochiscum var. tuberculatum 
G. M. Smith 

Euastrum abruptum f. minor 
West & West 

E. binale (Turp.) Ehrenberg 

E. ciastonii Raciborski 

E. gemmatum Brebisson 

Micrasterias Nordstedtiana, Wolle 







454 Rhodora 
Cosmarium Blytii Wille 

C. difficile var. dilatatwm Borge 

C. furcatospermum West & West 
C. granatum Brebisson 

C. isthmium W. West 

C. ornatum Ralfs 

C. portianum Archer 

C. pseudoconnatum Nordstedt 

C. pseudopyramidatum Lundell 

C. pyramidatum var. transitorium Heimer] 

C. tumidum Lundell 

Staurastrum dejectum Brebisson 

S. gracile var. nanum Wille 

S. gracile var. tenuissima Ralfs 

S. Leptocladum Nordstedt 

S. Ravenelii var. spinulosum Irenee-Marie 

Spondylosium planum (Wolle) 
West & West 

Desmidium Aptogonum Brebisson 

D. Baileyi (Ralfs) Nordstedt 

Xanthidium cristatum Brebisson 


Tabellaria fenestrata (Lyngb.) Kuetzing 
T. flocculosa (Roth) Kuetzing 

T. quadrisepta Knudson 

Meridion circulare (Grev.) Agardh 
Eunotia elegans Ost. 

E. formica Ehrenberg 

E. pectinalis (O.F.M.) Rabenhorst 

E. pectinalis var. ventricosa Grunow 

E. praerupta var. bidens (Ehr.) Grunow 
E. tenella (Grun.) Cleve 

E. serra var. diadema (Ehrenb.) Patrick 
Hannaea arcus Patrick 

Pinnularia flexuosa Cleve 

P. formica (Ehrenb.) Patrick 

Frustulia rhomboides (Ehrenb.) de Toni 




[Vol. 76 



1974] Algae on Haloragaceae — Colt & Hellquist 455 

E. vulgaris Thwaites IP-F-T 
Gomphonema acuminatum var. coronatum 

(Ehrenb.) W. Smith IP, PP-C-T 
G. augur Ehrenberg PP-C-T 


Microchaete tenera (Thuret) de Toni PP-F-T 
Hapalosiphon intricatus West & West IP-C-T 
Gloeotrichia Pisum (C. A. Agardh) 

Thuret IP-F-E 


ANDERSON, E. N., & E. R. WALKER. 1920. An ecological study of 
the algae of some sandhill lakes. Trans. Amer. Micros. Soc. 
39: 51-85. 

CHAPMAN, V. J. 1962. The Algae. MacMillan & Co., N. Y. 

Davis, C. C. 1955. The marine and fresh-water plankton. Mich. 
State Univ. Press, East Lansing, Mich. 

Fassett, NoRMAN C. 1966. A manual of aquatic plants. Univ. 
Wisc. Press, Madison, Wisc. 

FERNALD, M. L. 1950. Gray's Manual of Botany. American Book 
Co., New York, N. Y. 

IRENEE-MARIE, FRERE. 1939. Flore Desmidiale de la Region de 
Montreal. Laprairie, P. Q. 

JACKSON, DANIEL F. 1964. Algae and Man. D. F. Jackson, Ed. 
Plenum Press, New York, N. Y. 

NEEDHAM, JAMES G., & J. T. LLOYD. 1916. The life of inland waters. 
Comstock Publishing Co., Ithaca, N. Y. 

PATRICK, R., & C. W. REIMER. 1966. The Diatoms of the United 
States exclusive of Alaska and Hawaii. Monograph No. 13, 
Acad. Nat. Sci. Phila. 

Prescott, G. W. 1962. Algae of the Western Great Lakes Area. 
Wm. C. Brown Co., Dubuque, Iowa. 

1964. How to know the fresh-water Algae. Wm. C. 
Brown Co., Dubuque, Iowa. 

SMITH, G. M. 1924. Phytoplankton of the Inland Lakes of Wiscon- 
sin. Bull Univ. Wisc. Press, Serial 1048. 

1950. Fresh-water Algae of the United States. Mc- 
Graw-Hill Book Co., N. Y., N. Y. 

TIFFANY, L. H. 1951. Ecology of fresh water Algae. In: Manual 

of Phycology, Chronica Botanica, Waltham, Mass. 

BOSTON, MASS. 02115 

[Vol. 76 



Plate 1 

1974] Algae on Haloragaceae — Colt & Hellquist 457 

Plate 1: 

a) Eunotia formica Ehrenberg 

b) E. elegans Ost. 

c) Staurastrum leptocladum Nordstedt (semicell) 

d) Gomphonema acuminatum var. coronata (Ehr.) W. Smith 
e) G. auger Ehrenberg 

f) Spondylosium planum (Wolle) West & West 

g) Pinnularia formica, (Ehr.) Patrick 

h) Pediastrum duplex var. cohaerens Bohlin 

i) Desmidium Baileyi (Ralfs) Nordstedt 

j) Gloeotrichia Pisum (C. A. Agardh) Thuret 

k) Seytonema crispum (C. A. Agardh) Bornet 

1) Cosmarium isthmium W. West 

m) Tabellaria flocculosa (Roth) Kuetzing 

n) Eunotia praerupta var. bidens (Ehr.) Grunow 

o) Scenedesmus bijuga (Turp.) Lagerheim 

p) Characium ornithocephalum A. Braun 

q) Euastrum gemmatum Brebisson (semicell) 

r) Scenedesmus abundans var. brevicauda G. M. Smith 

458 Rhodora [Vol. 76 

"75w A, 50w J, "Sue bl, H24 C-i, km-p 

Plate 2 


Algae on Haloragaceae — Colt & Hellquist 459 

Plate 2: 




Oedogonium pseudoplenum Tiffany 

Oe. michiganense Tiffany 

Scenedesmus dimorphus (Turp.) Kuetzing 
Staurastrum dejectum Brebisson 

Eunotia serra var. diadema (Ehr.) Patrick 
Scenedesmus quadricauda var. maxima West & West 
Desmidium aptogomum Brebisson 

Staurastrum curvatum var. elongatum G. M. Smith 
Eunotia pectinalis (O.F.M.) Rabenhorst 
Pleurotaenium subcoronulatum var. detum West & West 
Euastrum abruptum fa. minor West & West 
Bulbochaete furberae Collins 

Pediastrum tetras var. tetraodon (Corda) Rabenhorst 
Euastrum ciastonii Rabenhorst 

Xanthidium cristatum Brebisson (semicell) 
Nephrocytium lunatum W. West 


Further field and herbarium studies of Illinois plants have 
added several new taxa to the state flora. Several are ad- 
ventive species from the Chicago area, while others repre- 
sent native species of taxonomically difficult groups and are 
treated in some detail. Appreciation is accorded to Mr. 
Floyd Swink and his colleagues at the Morton Arboretum 
who kindly made available some of their recent collections. 
All records cited are in the herbarium of Southern Illinois 
University (siU), the Morton Arboretum (MOR), or the 
Missouri Botanical Garden (MO). 

Poa bulbosa L. The bulbous blue grass, native to Eurasia, 
is previously unreported from Illinois. Collections have 
been made of this from cultivated beds at the Morton Ar- 
boretum. DU PAGE CO.: Morton Arboretum, within the shrub 
collection in Lacy Land, May 20, 1971, F. Swink (MOR). 

Phleum pratense L. f. viviparum (S. F. Gray) Louis- 
Marie. Vivipary, whereby plantlets or vegetative buds re- 
place the flowers in part or all of the inflorescence, may be 
encountered in several genera. The phenomonon seems most 
frequent in the Gramineae, especially such cool season 
grasses as Poa and Festuca. Vivipary in Scirpus atrovirens 
(Cyperaceae) is also commonly encountered. Because of 
the apparent rare occurrence of this type of apomictic re- 
production in Phleum pratense, this report seems worth- 
while. DU PAGE CO.: In the Morton Arboretum, July 27, 
1971, F. Swink (MOR). 

Fuirena scirpoidea Michx. The previously known range 
of this species was along the coastal plain of Georgia, 
Florida, Alabama, and Louisiana. Its discovery in Illinois 
at the Dolan Lake Conservation area marks a range exten- 
tion of more than 500 miles. One colony was found in 
shallow water growing beside Eleocharis obtusa. Svenson 


1974] Illinois Studies — Mohlenbrock & Evans 461 

(1957), the most recent monographer of the genus, com- 
bines F. pumila Torrey and F. breviseta Coville with F. 
squarrosa Michx. The three species now attributed to the 
Gray's Manual region may be identified in the following key. 

1. Leaves reduced to sheaths without blades. F. scirpoidea. 
1. Leaf blades well developed. ...................... 2. 
2. Achene dark brown with the beak pubescent; scale 
awn terminal. ............ ET . F. squarrosa. 

2. Achene yellow-brown with the beak 2 scale 
awn arising below the apex. ......... F. simplex. 

Although F. squarrosa (F. pumila) has been attributed 
to Illinois (Fernald, 1950; Jones 1950, 1963; Patterson, 
1876), a thorough search by the senior author has failed 
to reveal any such specimens from this state. HAMILTON 
CO. : Shallow water, Dolan Lake shore in Dolan Conservation 
area, June 13, 1970, N. Tracy (SIU). 

Salix X myricoides Muhl. This taxon is a reputed hybrid 
between S. rigida Muhl. and S. sericea Marsh., although it 
appears to have more of the characteristics of S. sericea. 
It differs from S. sericea by its canescent twigs and thinly 
sericeous capsules, while S. sericea has glabrous or glabrate 
twigs and densely sericeous capsules. We have recently 
collected it from Jackson County for its first known locality 
in Illinois, but it undoubtedly occurs elsewhere in the state. 
JACKSON CO.: Along rocky stream, Giant City State Park, 
August 30, 1971, R. H. Mohlenbrock (SIU). 

Populus X smithii Boivin. This is the hybrid between 
P. grandidentata Michx. and P. tremuloides Michx., first 
described from Michigan by Wagner in 1970. It is inter- 
mediate in several characters between the parent species, 
the most obvious being the number of dentations along the 
leaf margins. The following key will serve to distinguish 
this hybrid from its parents. 

1. Margin of leaf dentate, with 5-25 teeth (averaging 10- 
20)? budà pubescent. ............... ce wee ces 2. 

462 Rhodora [Vol. 76 

2. Margin of leaf with 5-15 teeth (averaging 10); 
petiole 5-10 em long (averaging 7 em). .......... 
SNNT? P. grandidentata. 

2. Margin of leaf with 12-25 teeth (averaging 20); 
petiole 3-6 cm long (averaging 5.5 em). .......... 
EEDUVUOOÓO nul P. X smithü. 

1. Margin of leaf finely crenate, with 20 or more teeth 

(averaging 31) ; buds glabrous or nearly so. ...... ... 

VN P. tremuloides. 

LA SALLE CO.: Starved Rock State Park, June 16, 1943, 
G. N. Jones 15791 (MO). PEORIA CO.: Horse Shoe Bottom, 
July 27, 1919, V. H. Chase 3217 (Mo). 

Carya pallida (Ashe) Engl. & Graebn. Correspondence 
with Dr. Wayne Manning, noted authority on the genus 
Carya, suggested the possibility of C. pallida in Illinois, 
since it occurs in western Kentucky and southern Indiana. 
Consequently, the senior author has kept a close vigil for 
this species in southern Illinois and was rewarded with the 
discovery of a specimen from Union County referable to 
C. pallida. The distinction between C. pallida and C. texana 
Buckl. is difficult. Both have yellow-lepidote buds, and the 
fruits are not different, although the fruits in C. pallida are 
usually smaller. Carya pallida usually has at least a few 
leaves with tomentose rachises, while C. texana usually does 
not except in var. villosa (Sarg.) Little. In the spring, C. 
texana has distinctive red hairs, while C. pallida does not. 
UNION CO.: Dry, acidic woods, along Grapevine Trail, May 
21, 1972, R. H. Mohlenbrock (SIU). 

Quercus veiutina Lam. f. missouriensis (Sarg.) Trel. 
This form of Black Oak has the lower surface of the leaves 
densely and permanently pubescent. In this respect, f. mis- 
souriensis resembles Q. falcata Michx., but differs from this 
species in leaf shape and bud and acorn characteristics. 
SALINE CO.: Edge of dry woods, Old Stone Face, July 3, 
1971, R. H. Mohlenbrock (SIU). 

Chenopodium. While preparing the treatment of Cheno- 

1974] Illinois Studies — Mohlenbrock & Evans 463 

podium for The Illustrated Flora of Ilinois series, the 
senior author has studied many collections of the genus 
and has re-evaluated the species previously known from the 
state. He has followed closely the treatment by Steyermark 
(1963) in Flora of Missouri in order to make more uniform 
the treatment of this genus in the central midwest. Three 
additional recognizable species from Illinois emerged from 
the study. 

Chenopodium pallescens Standl. In general, all Illinois 
Chenopodium with white-mealy, narrow, entire leaves have 
been designated either C. leptophyllum Nutt. (Fernald, 
1950; Gleason, 1952) or C. pratericola Rydb. (Jones, 1963). 
Illinois material can be separated reliably into two taxa, 
one to be called C. pallescens Standl, the other C. desic- 
catum A. Nels. var. leptophylloides (Murr.) Wahl. The 
following key serves to separate these taxa. 

1. Leaves 1-nerved, linear; pericarp firmly attached to 

readily removed from seed. ©... 1 eee eee nn nn 
C. desiccatum var. leptophylloides. 

Chenopodium pallescens is found in rocky ground and in 
waste areas, primarily in the north-central counties of the 
state. Chenopodium dissectum var. leptophylloides is almost 
always in waste ground in the northern half of Illinois. 

Chenopodium bushianum Aellen. Although Jones (1963) 
combined this species with C. album L., the conspicuous 
honeycombed surface of the seeds of C. bushianum makes 
it a readily recognizable species. It is found occasionally 
throughout the state in fields, woods, and waste places. 

Chenopodium strictum Roth var. glaucophyllum (Aellen) 
Wahl. This taxon is related to C. album L. and C. mis- 
souriense Aellen. It differs from C. album by its smaller 
seeds (0.8-1.2 mm broad), and from C. missouriense by its 
coarsely toothed lower leaves and its calyx lobes which do 

464 Rhodora [Vol. 76 

not cover the fruit. JACKSON CO.: Waste ground, one mile 
north of Carbondale, September 3, 1971, R. H. Mohlenbrock 

Draba verna L. var. boerhaavii Van Hall. This variety 
differs from typical var. verna by its fruits that are never 
twice as long as broad and by the seeds fewer than 40 per 
fruit. It is naturalized from Europe and adventive in a 
mowed field in Illinois. JACKSON CO.: Field, Southern Illi- 
nois University campus, Carbondale, April 4, 1972, R. H. 
Mohlenbrock (SIU). 

Draba reptans (Lam.) Fern. var. micrantha (Nutt.) Fern. 
This variety, unreported previously from Illinois, occurs 
occasionally in northern Illinois where it sometimes is found 
growing with var. reptans. Variety micrantha differs by 
its hispidulous fruits. 

Suaeda depressa (Pursh) S. Wats. This species is nor- 
mally found in the naturally occurring saline soils of south- 
ern Canada, through the plains states, and south to Texas. 
In an apparent response to salting by the highway depart- 
ment, the plant grew along the highway median in the 
Chicago area. COOK CO.: Median strip, one mile north of 
Illinois 22, one mile south of Everett Pond, October 7, 1972, 
R. Schulenberg (MOR). 

Spergu'aria media (L.) C. Presl. Another adventive ele- 
ment new to the Illinois flora is this member of the pink 
family. The previous range of this species appears to be 
the natural saline soils of central and coastal New York as 
well as coastal California and Oregon. Illinois collections 
were made near Elgin along a tollway where salt is fre- 
quently used by the highway department. KANE CO.: 314 
miles west of the Elgin toll booth on the Northwest Tollway 
right-of-way bank, in sterile clay, September 3, 1972, R. 
Read. (MOR). 

Pyrus pyrifolia (Burm. f.) Nakai. The Chinese Pear is an 
occasional cultivated plant in Illinois. A specimen about 

1974] Illinois Studies — Mohlenbrock & Evans 465 

thirty-five feet tall has been found growing along a rocky 
stream in Giant City State Park. This species differs from 
the more common P. communis L. by its rounder fruits, 
slightly larger flowers, and sharply toothed leaves. JACKSON 
CO.: Along Stonefort Creek, Giant City State Park, Septem- 
ber 11, 1970, R. H. Mohlenbrock (SIU). 

Crataegus marshallii Egglest. This is one of the few spe- 
cies of Crataegus in Illinois which has the veins of the leaf 
running to the sinuses as well as to the points of the lobes. 
It differs from C. phaenopyrum (L. f.) Medic. by its petioles 
much longer in relation to the length of the blade, its fewer 
stamens with reddish anthers, and by its usually two nut- 
lets per fruit. Its habitat in swampy woods is also dis- 
tinctive. JACKSON CO.: Swampy woods, Greentree Reser- 
voir, 2144 miles southwest of Gorham, R. Anderson (SIU). 

Triadenum virginicum (L.) Raf. The discovery of this 
species in Lake County brings to four the number of spe- 
cies of Triadenum known from Illinois. The following key 
separates these four species. 

1. Leaves without punctations. .  ....... T. tubulosum. 
1. Leaves punctate, at least on the lower surface. .... . 2. 
2. Leaves petiolate. ................ ... T. walteri. 

2. Leaves sessile. ....... DEMNM o re 9. 

3. Sepals obtuse, up to 5 mm long; styles up to 1.5 

mm Jong, ........... aa REA T. fraseri. 

3. Sepals acute, 5-8 mm long ; styles 2-3 mm long. 
E. laeua gy ax v7 12 JE NEN T. virginicum. 

LAKE CO.: Boggy ground, Illinois Beach State Park, Au- 
gust 3, 1972, R. H. Mohlenbrock (SIU). 

Mentzelia decapetala (Pursh) Urban & Gilg. This species 
is native to dry prairies and plains west of the Mississippi 
River. In Grundy county, apparently its easternmost site, 
it was collected along a railway where it occurred with 
other such adventives as Kochia scoparia, Setaria viridis, 
Helianthus annuus, and Grindelia lanceolata f. latifolia. 

466 Rhodora [Vol. 76 

GRUNDY CO.: Cinder and gravel ballast, at Eileen, along 
right-of-way of Santa Fe RR, August 13, 1972, R. Schulen- 
berg, D. Kropp, & D. Ladd (MOR). 

Polypremum procumbens L. This species was expected to 
occur in southern Illinois since it is known from Kentucky 
to the south and five counties in nearby southeastern Mis- 
souri. Collections were made from a sandy field in the 
Horseshoe Lake Conservation area. The inconspicuous 
corolla, procumbent habit, and slender leaves probably have 
resulted in this species being overlooked in other localities. 
ALEXANDER CO.: Low, sandy field, Horseshoe Lake, August 
11, 1971, J. Huston 920 (SIU). 

Jacquemontia tamnifolia (L.) Griseb. This species is na- 
tive to the southeastern United States and represents a 
genus new to the Illinois flora. This probable adventive was 
collected from a disturbed habitat where it grew in associa- 
tion with Agropyron repens, Ipomoea purpurea, and Eu- 
phorbia dentata, The following key serves to distinguish 
Jacquemontia from Ipomoea, Convolvulus, and Calystegia in 

1. Stigma 1, capitate or with 2 to 3 lobes. ...... Ipomoea. 
1. Stigmas 2, noncapitate, without lobes. ............ 2. 
2. Stigmas elliptic, oblong or flattened. . Jacquemontia. 

2. Stigmas filiform or subulate. ... ........... . 9. 

3. Calyx not concealed by large bracts; fruit 2-locu- 

lar. ....... see Convolvulus. 

3. Calyx concealed by 2 large bracts; fruit 1-locular. 
BENNETT UU Calystegia. 

GRUNDY CO.: Railroad tracks at Gardner, October 15, 1972, 
R. Schulenberg & E. Lace (MOR). 

Vinca major L. The large-flowered periwinkle, an intro- 
duced European cultivar, was previously known in the wild 
in the coastal states from North Carolina south to Missis- 
sippi. In Illinois the species was collected around an old 
homestead in Pope County where it grew without cultiva- 
tion. POPE CO.: Randolph Farm, V. Randolph (SIU). 

1974] Illinois Studies — Mohlenbrock & Evans 467 

Matelea decipiens (Alex.) Woodson. This climbing mem- 
ber of the milkweed family is widely distributed in central 
and southern Missouri, but the record cited below is ap- 
parently the first from Illinois. All three Illinois repre- 
sentatives of this genus are restricted to the southernmost 
counties. They may be distinguished by the following key. 

1. Flowers greenish-yellow; pedicels glabrous; follicles 
smooth, angular. ......... 0 .........., M. gonocarpa. 
1. Flowers rose, maroon, or rarely cream; pedicels pubes- 

2. Petals 1.5-2.5 mm wide, rose or rarely cream. ..... 
CETERAE id en vue 39 oa A RE yo G's UR tw deen M. obliqua. 

WILLIAMSON CO.: Low floodplain woods, natural levee of 
Big Muddy River, north of Colp, May 28, 1972, M. Swayne 
& J. Swayne (SIU). 

Physalis macrophysa Rydb. This plant, treated variously 
as a species, variety, or form, differs from P. subglabrata 
Mack. & Bush by its translucent leaves and its large, pyra- 
midal calyces when in fruit. The calyx becomes 3-6 cm 
long during fruiting. Illinois specimens, previously labelled 
P. subglabrata, have been seen from Champaign and Peoria 

Antirrhinum orontium L, The often cultivated lesser 
snapdragon has been collected from a cultivated oat field. 
Although it has probably escaped elsewhere in Illinois, this 
is the first report of such an occurrence. DU PAGE CO.: Oat 
field, National Accelerator Laboratory near Warrenville, 
September 28, 1972, F. Swink (MOR). 

Penstemon cobaea Nutt, This sometimes cultivated 
beards-tongue was previously known from southern Mis- 
souri northwest to Nebraska, south to Texas, and east to 
Arkansas. Within this range the typical habitats are prai- 
ries, glades, and bluffs. In Illinois this species was collected 
from a dry meadow where it grew in association with such 

468 Rhodora [Vol. 76 

prairie elements as Asclepias verticillata, Poa compressa, 
and Physalis subglabrata. Although this species is not 
known to be native to northeastern Illinois, the plant asso- 
ciates as well as the prairie habitat might suggest an indig- 
enous condition. KANE CO.: Flat, dry meadow, west of 
Montgomery, near Kendall County line, June 25, 1972, J. 
Phillips, D. Young, & R. Schulenberg (MOR). 

Lagenaria siceraria Standl. This highly variable gourd is 
frequently planted in Illinois and occasionally escapes from 
cultivation. An early collection from Hancock County and 
a recent one from Jackson County verify its existence in 

Cucurbita pepo L. var. ovifera (L.) Alef. This variety of 
the common Field Pumpkin has extremely variable fruits, 
many of them the source of interesting ornamental gourds 
such as the Pear, the Bell, the Apple, the Egg, and the 
Orange. A single collection has been made for this plant in 
Illinois. UNION CO. : in disturbed soil along a stream, 2 miles 
north of McClure, July 31, 1971, R. H. Mohlenbrock (SIU). 

Sherardia arvensis L. This new element of the Illinois 
flora is adventive but well established in areas of the SIU 
campus at Carbondale. It may be confused with Galiwm 
but is distinguished by its long, funnel-form, blue corolla 
and narrow leaf-like involucres that subtend the flowers. In 
addition to our location, the species is local in Nova Scotia, 
southeastern Canada, Missouri, Tennessee, and North Caro- 
lina. JACKSON CO.: Lawn, SIU campus by the Forest Ser- 
vice Laboratory, April 30, 1971, D. Evans 11408 (SIU). 

Campanula rotundifolia L. var. velutina A. DC. The oc- 
currence of typical C. rotundifolia from the rocky environs 
of northeastern Illinois is not uncommon. However, variety 
velutina is heretofore reported only from a single collection 
by E. J. Hill from Cheboygan County in northern Michigan. 
We have this taxon from near Apple River Canyon State 
Park. Here the plant grows in the crevices of an east-facing 
limestone wall which borders Apple River. Associated spe- 

1974] Illinois Studies — Mohlenbrock & Evans 469 

cies include the typical form, Aquilegia canadensis, Carex 
granularis, and the cliff fern Cryptogramma stelleri, Con- 
sidering the close habitat association with the markedly 
different typical form, it seems unlikely that this pubescent 
variety is a mere ecological variant as claimed by some. 
JO DAVIESS CO.: East-facing limestone wall bordering Apple 
River, 14 mile northeast of Apple River Canyon State Park, 
June 13, 1972, D. Evans 1156 (SIU). 

Grindelia lanceolata Nutt, f. latifolia Steyerm. Prior to 
this report from Grundy County, f. latifolia was unknown 
in Illinois. The typical form remains unreported from the 
state. The closest occurrence of this new taxon to the Il- 
linois site is the type locality in Stone County, Missouri. 
Collections are also known from Oklahoma. GRUNDY CO.: 
At Eileen, in N 14 of Sect. 36, T33N, R6E, August 13, 1972, 
R. Schulenberg & D. Ladd (MOR). 

Senecio jacobaea L. Collections from railways and road- 
sides often produce rare or unusual adventive species. This 
waif recently has been collected from a railway in the Chi- 
cago area. DU PAGE CO.: Great Western RR at Euclid Ave- 
nue, Villa Park, August 17, 1972, F. Swink (MOR). 

Thelesperma gracile (Torr.) Gray. The natura] habitat 
and range of this unusual composite are the prairies and 
plains from Nebraska to Wyoming, south to Arizona, and 
east to Texas and Oklahoma; also Mexico and South Amer- 
ica. In addition, Steyermark (1963) reports two adventive 
sites in Missouri. This roadside collection from Kane county 
represents the easternmost extension of its range in North 
America. KANE CO.: Along Ancutt Road, west of Montgom- 
ery, July 4, 1972, F. Swink (MOR). 


FERNALD, M. L. 1950. Gray's Manual of Botany, ed. 8. The Ameri- 
can Book Company, New York. 1632 pp. 

GLEASON, H. A. 1952. The New Britton and Brown Illustrated 
Flora of Northeastern United States and Adjacent Canada. New 
York Botanical Garden, New York. 3 Vol. 1762 pp. 

470 Rhodora [Vol. 76 

Jones, G. N. 1950. Flora of Illinois, ed. 2. Am. Midl. Nat. Monogr. 

5: 1-368. University of Notre Dame Press, Notre Dame, Indiana. 
1963. Flora of Illinois, ed. 3. Am. Midl. Nat. Monogr. 

7: 1-401. University of Notre Dame Press, Notre Dame, Indiana. 

PATTERSON, H. N. 1867. Catalogue of phaenogamous and vascular 
eryptogamous plants of Illinois. Oquawka, Illinois. 54 pp. 

STEYERMARK, J. A. 1963. Flora of Missouri. The Iowa State Uni- 
versity Press, Ames, Iowa. 1725 pp. 

SvENSON, H. K. 1957. Fuirena. In: North American Flora 18(9): 



While collecting in the Ocean Park section of Old Orchard 
Beach, York County, Maine, in recent summers, I have 
noticed certain plants of Spiraea latifolia, Meadowsweet, 
with deep-pink flowers. Some of these plants have pubes- 
cent panicles; others are glabrous. I have found this form 
growing both in open wet spots and in shaded woodland 

Spiraea latifolia (Ait.) Borkh. forma rosea T. W. Wells, 
forma nova. TYPE: Maine: YORK co.: Old Orchard Beach, 
11 October, 1969, T. W. WELLS (NEBC). Differt a forma 
latifolia floribus valde roseis. 

While collecting in salt-marshes along Goose Fare Creek, 
which is the southern boundary of Ocean Park, I found 
plants of Polygonum ramosissimum which differ from the 
typical in that the sepals have bright-white margins. 

Polygonum ramosissimum Michx. forma alba T. W. Wells, 
forma nova. TYPE: Maine: YORK co.: Old Orchard Beach, 
5 September, 1970 T. W. WELLS (NEBC). Differt a forma 
ramosissimum sepalis albomarginatis. 




In recent years the flora of the Bahama Islands has again 
aroused the interest of several taxonomists, who have found 
The Bahama Flora of N. L. Britton and C. F. Millspaugh 
(1920) to be inadequate in many respects. Not only have 
numerous nomenclatural changes been proposed, but also 
since that time collections of species new to the islands have 
been made (Howard, 1950; Howard & Dunbar, 1964; Lewis, 
1971). At the present time The Fairchild Tropical Garden 
is undertaking the much-needed revision of the outdated 

This writer has made two collecting trips to the Bahamas; 
the first from April 28 to May 16, 1970, on New Providence 
Island, Long Island, and Inagua Island, and the second from 
May 18 to June 3, 1972, on Long Island alone. Both trips 
were as an undergraduate at Bates College, Lewiston, 
Maine, and were under the guidance and direction of Dr. 
Harold E. Hackett of the Department of Biology at that 
institution. Four collections were from New Providence, 
147 from Long Island, and 51 from Inagua. It was noted 
that the flora of Long Island, a flora which seems to re- 
semble that of the eastern coast of Cuba, was particularly 
poorly known. Long Island is an “out island" nearly one- 
hundred miles long and four miles wide, trending north- 
south, and bisected by the Tropic of Cancer. Since the flora 
was poorly known, Walter Cerbin, SSJ, of Clarence Town, 
Long Island agreed to collect in his area, with the process- 
ing of specimens to be done at the New York Botanical 
Garden. To date Father Cerbin's collections number 76 (nos. 
102-167 from Long Island, and 1001-1010 from Inagua). 
These are deposited in this writer's personal herbarium 
(designated as SRH) housed at the New York Botanical 
Garden, with duplicates at New York and the Arnold Ar- 


472 Rhodora [Vol. 76 

According to Britton and Millspaugh (1920), five col- 
lectors had gathered specimens on Long Island up to the 
time of their work, and of their collections only 173 num- 
bers were cited (nos. 495-520, 524 of William C. Coker, 
1903; and nos. 6225-6307 of Britton and Millspaugh, 1907). 
The two others considered to have collected on the island 
were Swainson (1830-42), whose collections were not known 
to the authors of The Bahama Flora, and H. F. A. von 
Eggers (1888), whose collections were apparently not suf- 
ficiently well-known to be cited. Among those who have 
touched upon Long Island since that time are Dr. Harold 
E. Hackett of Bates College (53 numbers in the Duke Uni- 
versity Herbarium), Father Walter Cerbin, and this writer. 

The results of the recent collections include several ncw 
records: 49 species newly reported for Long Island, 4 new 
to Inagua, and 8 new to the Bahama Islands. The collections 
also include the rediscovery of a species formerly known 
only from the type, Zamia lucayana Britton. It is interest- 
ing to note that many of these new records are of rather 
common species in the West Indies, reflecting the paucity 
of the collections from the area to date. Cultivated plants 
have been included since they commonly escape on the 
islands and become a part of the local flora. 

The list of taxa is divided into four sections: 1) species 
newly reported for Long Island, 2) species newly reported 
for Inagua, 3) species newly reported for the Bahamas, and 
4) a rediscovery. The citations after the collection number 
refer to those institutions in which the specimen is housed, 
and the abbreviations are as in the Index Herbariorum 
(Lanjouw and Stafleu, 1964). 

The author wishes to thank Dr. William T. Gillis for his 
help in reviewing the manuscript, Dr. Harold E. Hackett 
for his guidance and encouragement while in the Bahamas 
and at Bates College, Drs. Arthur and Noel Holmgren, Dr. 
Gerrit Davidse, and Mr. Kenneth Becker for their help with 
certain determinations, and Dr. John T. Mickel, his advisor 
at the New York Botanical Garden. 

1974] Bahama Flora — Hill 473 

I. Species Newly Reported for Long Island. 
A. Native or long established species. 
POLYPODIACEAE: Acrostichum danaeifolium Langsd. 
& Fisch.: Hill 840 (sRH) sinkhole, Village Road, 
S. Clarence Town. 

Asplenium dentatum L.: Hill 834 (A, SRH) sink- 
hole, Deadman’s Cay. 

Pteris longifolia L.: Hill 819b (SRH) moist de- 
pression, Mangrove Bush. 

Thelypteris normalis (C. Chr.) Moxley: Hill 125, 
835 (SRH) sinkholes, Village Road, S. Clarence 

GRAMINEAE: Chloris petraea Sw.: Hill 467 (SRH) 
Clarence Town. Det. G. Davidse. 

Distichlis spicata (L.) Greene: Hill 123 (A, SRH) 
Harbor Point, Clarence Town. Det. A. Holmgren. 

Uniola paniculata L.: Hill 826 (A, SRH) Harbor 
Point, Clarence Town. 

AMARYLLIDACEAE: Agave sisalama (Engelm.) Per- 
rine: Hill 824 (sRH) Clarence Town. 

Hymenocallis declinata (Jacq.) M. Roem.: Hill 
829 (A, SRH) Turtle Cove. 

BROMELIACEAE: Tillandsia usneoides L.: Hill 670b 
(US, A, SRH) Galloway's Cave, South Clarence Town. 

Tillandsia utriculata L.: Hill 529 (sRH) Village 
Rd., S. Clarence Town. 

COMMELINACEAE: Rhoeo spathacea (Sw.) Stearn: 
Hill 244 (sRH) Clarence Town. 

ORCHIDACEAE: Encyclia bahamensis (Griseb.) Brit- 
ton: Hill 532 (US, A, SRH) 4 mi. W. of Clarence 

POLYGONACEAE: Antigonum leptopus H. & A.: Cer- 
bin 120 (NY, A, SRH) Clarence Town. 

Coccoloba swartzii Meisner: Hill 471, 841 (SRH) 
Clarence Town. 

NYCTAGINACEAE: Commicarpus scandens — (L.) 
Standley: Cerbin 104 (NY, A, SRH) Turtle Cove. 


Rhodora [Vol. 76 

Guapira bracei (Britt.) Little: Hill 848 (SRH) 
Turtle Cove. 
BATIDACEAE: Batis maritima L.: Hill 531 (SRH) 
Clarence Town. 
MIMOSACEAE: Acacia farnesiana (L.) Willd. : Cerbin 
144 (NY, A, SRH) Village Road, South Clarence 

Albizia lebbeck (L.) Benth.: Hill 540 (SRH) 
Clarence Town. 
CAESALPINIACEAE: Cassia biflora L.: Cerbin 116 
(NY, A, SRH). 
ZYGOPHYLLACEAE: Kallstroemia maxima (L.) T. & 
G.: Hill 952 (SRH) Clarence Town. 
EUPHORBIACEAE: Pedilanthus tithymaloides (L.) 
Poit.: Hill 139 (SRH); Cerbin 139 (NY, A, SRH) 
Clarence Town. 

Ricinus communis L.: Hill 542 (SRH) common 
escape in Clarence Town. 
MALVACEAE: Abutilon trisulcatum (Jacq.) Urban: 
Cerbin 160 (NY, A, SRH) Clarence Town. 

Cienfuegosia yucatanensis Millspaugh: Hill 1015 
(SRH) moist depression between Hamilton's and 
Buckley's. Identified by Dr. Paul Fryxell. 

Thespesia populnea (L.) Soland.: Hill 129 (a, 
SRH) Clarence Town. 
TURNERACEAE: Turnera diffusa Willd.: Hill 845 
(SRH) Harbor Point, Clarence Town. 
PASSIFLORACEAE: Passiflora bahamensis Britton: 
Hill 515b (A, SRH); Cerbin 166 (NY, A, SRH) Clar- 
ence Town. 
CARICACEAE: Carica papaya L.: Hill 142 (A, SRH) 
occasional in Clarence Town. 
SAPOTACEAE: Mastichodendron foetidissimum 
(Jaeq.) Cronq.: Hill 839 (A, sRH) Turtle Cove. 
APOCYNACEAE: Plumeria obtusa L.: Hill 831 (SRH) 
Turtle Cove. 
ASCLEPIADACEAE: Cryptostegia grandiflora (Roxb.) 


Bahama Flora — Hill 475 

R. Br.: Hill 128 (SRH); Cerbin 153 (NY, SRH) 
Clarence Town outskirts. 
CONVOLVULACEAE: Evolvulus squamosus Britton: 
Cerbin 113 (NY, A, SRH) Mangrove Bush. 

Merremia dissecta (Jacq.) Hall: Cerbin 117 (Ny, 
A, SRH) Morrisville. 
SCROPHULARIACEAE: Stemodia maritima L.: Hill 947 
(SRH); Cerbin 119 (NY, A, SRH) 4 miles west of 
Clarence Town. Determined by Dr. Noel Holmgren. 
ACANTHACEAE: Blechum brownei Juss.: Cerbin 126 
(A, NY, SRH) Clements. 

Ruellia tuberosa L.: Hill 846 (SRH) Clarence 
RUBIACEAE: Casasia clustifolia (Jacq.) Hitch.: Cer- 
bin 141, 152 (NY, A, SRH) Salt Pond. 
ASTERACEAE: Bidens pilosa L.: Cerbin 111 (NY, A, 
SRH) Mangrove Bush. 

Wedelia trilobata (L.) Hitche.: Cerbin 136 (Ny, 
A, SRH) Salt Pond. 

Verbesina encelioides (Cav.) Benth. & Hook.: 
Cerbin 159 (NY, A, SRH) Clarence Town. Deter- 
mined by K. Becker. 

B. Cultivated or escaped. 

AMARYLLIDACEAE: Hippeastrum puniceum (Lam.) 
Urban: Hill 1017 (sRH) Hamiltons. 

FABACEAE: Sesbania grandiflora (L.) Pers.: Cerbin 
102 (NY, A, SRH) Turtle Cove. 

EUPHORBIACEAE: Codiaeum variegatum (L.) Blume: 
Cerbin 122 (NY, A, SRH) Clarence Town. 
ARALIACEAE: Polyscias guilfoylei (Bull. Bailey: 
Cerbin 114 (NY, A, SRH) Morrisville. 
APOCYNACEAE: Thevetia peruviana (Pers.) K. 
Schum.: Hill (A, SRH) Clarence Town. 

OLEACEAE: Jasminum fluminense Vell.: Cerbin 140 
(NY, A, SRH) Salt Pond. 

SCROPHULARIACEAE: Russellia equisetiformis Schl. 
& Cham.: Cerbin 133 (NY, A, SRH) Deadman’s Cay. 




Rhodora [Vol. 76 

CUCURBITACEAE: Momordica charantia L.: Cerbin 
125 (NY, A, SRH) Clements. 

Species Newly Reported for Inagua. 

LILIACEAE : Sansevieria hyacinthoides (L.) Druce: Cer- 
bin 1001 (NY, A, SRH) Along road to Northwest Point. 
PASSIFLORACEAE: Passiflora bahamensis Britton: Hill 
515 (sRH) Union Creek. 

APOCYNACEAE: Nerium oleander L.: Hill 488a & b. 
(SRH) Matthew Town. 

CONVOLVULACEAE: Merremia dissecta (Jacq.) Hall: Hill 
489 (SRH) Matthew Town. 

Species Newly Reported for the Bahama Islands. 
AMARYLLIDACEAE: Crinum asiaticum L.: Cerbin 124 
(NY, A, SRH) Clarence Town, Long Island. Cultivated 
and possibly escaped in moist places. 

Zephyranthes citrina Baker: Hill 827 (A, SRH) Clar- 
ence Town, Long Island. 

Zephyranthes puertoricensis Traub: Hill 836 (SRH) 
Hamilton’s, Long Island. Also in cultivation from this 
population at NY. 

AMARANTHACEAE: Alternanthera brasiliana  (L.) 
Kuntze: Cerbin 127 (NY, A, SRH) Clements, Long Is- 
land. Determined by Mr. Kenneth Becker, NYBG. 
CAESALPINIACEAE: Bauhinia variegata L.: Hill 566 
(SRH) Nassau, New Providence. Commonly cultivated 
and escaped. 

MALPIGHIACEAE: Malpighia glabra L.: Cerbin 1005 
(NY, A, SRH) Matthew Town, Inagua. Cult. Cultivated 
and possibly escaped. 

CACTACEAE: Opuntia rubescens (Salm-Dyck) Lemains: 
Hill 476 (sRH) Matthew Town, Inagua. Dooryard. 
SOLANACEAE: Datura inoxia Mill.: Hill 850 (SRH) Clar- 
ence Town, Long Island. Roadside. Also Nash and 
Taylor 1357, formerly determined as D. metel L. but 
re-identified by H. N. Moldenke. Inagua. 

IV. Rediscovery. 

CYCADACEAE: Zamia lucayana Britton: Hill 828 (Mo, 

1974] Bahama Flora — Hill 477 

K, US, A, F, BM, TRIN, SRH, NY). Found in limestone and 
sand 100 yards from the sea, about five miles north of 
the type locality; Turtle Cove, Long Island. Formerly 
known only from the type collection “In a sandy coastal 
thicket Clarence Town, Long Island, only one plant 
found after a long search (Britton and Millspaugh 
6271)” (Britton, 1907). This writer discovered an 
extensive colony with many old plants as well as a 
thriving understory of seedlings and hermit crabs 
which were feeding on the fleshy seed coats. Plants 
in the shade generally exhibited much longer leaves 
than those in sun, those specimens in shade agreeing 
more closely with the type. Male and female cones 
were collected, and living material is in cultivation at 
NYBG, Harvard, and the Fairchild Tropical Garden. 


BRITTON, N. L. 1907. Contributions to the Flora of the Bahama 
Islands. IV. Bull. N. Y. Bot. Gard. 5: 311. 

BRITTON, N. L. & C. F. MiLLsPAUGH. 1920. The Bahama Flora. 
Hafner Publishing Co., Inc. New York, 

Howanp, R A. 1950. Vegetation of the Bimini Island Group. Ecol. 
Monog. 20: 317-349. 

, & H. F. DUNBAR. 1964. Additions to the Flora of 

Inagua, the Bahamas. Rhodora 66: 6-15. 

LANJOUW, J. & F. A. STAFLEU. 1964. Index Herbariorum. 5th. 
ed. Part I. The Herbaria of the World. Reg. Veg. 31. Utrecht. 

LEWIS, W. H. 1971. Additions to the Flora of the Bahama Islands. 
Rhodora 73: 46-50. 




A dozen years ago a strikingly different Lechea was 
noted (Rhodora 63: 117. 1961.) among collections from 
southern Florida. It then seemed unwise, however, to de- 
scribe the population that they represent as a distinct new 
taxon since the specimens available were not as complete 
or mature as would be desired in proposing a new entity in 
such a perplexing and technical genus. However, additional 
collections from southern Florida and the low-keyed but 
persistent hints from Dr. Olga Lakela that something 
wasn’t quite satisfactory with available treatments finally 
prodded me into taking another look. Her own collections 
have largely eliminated one of my reservations about for- 
mally proposing a new taxon, although additional collections 
from throughout the growing season are still very much to 
be desired. Practically nothing is known of the plants ex- 
cept in the fruiting stage and although that is the most 
important stage for the accurate identification of members 
of this genus, it still would be most desirable to know 
something more about the vegetative features of this spe- 
cies. Leaves in this species are almost completely lacking 
by the time fruits are ripening. Still it seems probable that 
more of a stimulus for additional collections of this taxon 
and of the other highly distinctive representatives of the 
genus in peninsular Florida will be forthcoming if the taxon 
is formally proposed now than if we wait for more col- 


1. Pubescence of aerial stems mostly strongly divergent 
and spreading. ......ssse e nn 2. 

"This study was supported by NSF GB-13815, for which grateful 
acknowledgment is made. 



Genus Lechea — Wilbur 479 

2. Internal sepals conspicuously V- or U-shaped in 

cross-section with thin scarious margins and a 
roughened, often sparsely pilose keel (the sepal 
otherwise glabrous) ; leaves often over 1.5 cm. long; 
capsule thin-walled, readily splitting into 3 valves 
at maturity, subglobose, about equaling the calyx in 
length. .... 1. L. mucronata Raf. (L. villosa Ell.). 

Internal sepals but slightly bowed in cross-section 
with texture appearing uniform and pubescence scat- 
tered across surface; leaves all less than 1 cm. long; 
capsule thick-walled, indehiscent, ellipsoid or some- 
what rounded- barrell-shaped, exserted from the 
calyx for at least 1/3-1/2 its length. .............. 
ems Lis 2. L. divaricata Shuttlew. ex Britt. 

1. Pubescence of aerial stems mostly closely appressed or 
THOME ea iio ae on an a e oo owe ees ale ee 3. 


External sepals equaling or exceeding the internal 
sepals in length. ........... 00. ce eee n n n 4. 

4. External sepals at least one-fifth longer than the 
inner sepals and usually equaling or exceeding 
the capsule in length; capsule equaling or exceed- 
ing the inner sepals by not more than one-fifth its 
length; cauline leaves usually elliptic to elliptic- 
oblong, usually less than 5 times as long as wide, 
those below the inflorescence commonly appear- 
ing whorled and often more than 2 mm. wide. 
ck ct acs eo tie eet zu. De dA minor L. 

4. External sepals about equaling the inner sepals 
and never equalling the capsule in length; capsule 
usually exceeding the inner sepals by about 1/3- 
1/2 its length; cauline leaves narrowly oblong to 
linear, usually 6 times (or more) as long as wide, 
those below the inflorescent alternate and less 
than 2 mm. wide. ..........-..- rnm n 
HERR 4. L. sessiliflora Raf. (L. patula Legg.). 

3. External sepals shorter than the internal sepals. . 5. 

5. Leaves pubescent on both upper and lower sur- 


Rhodora [Vol. 76 

faces (at least those of the basal shoots conspicu- 
ously pilose above and below while the cauline 
and rameal leaves are usually inconspicuously 
pubescent over entire surface); flowers or fruits 
mostly clustered in 2's or 3's; capsule wall thick- 
ened and indurate. ........ 5. L. cernua Small. 

Leaves variously pubescent below but glabrous on 
upper surface; flowers or fruits not appearing 
fascicled (but obviously attached separately) ; 
capsule wall thin or at least not conspicuously 
indurate. e 6. 
6. Aerial stems perennial, suffruticose, clearly 
woody at base, with wiry woody branches; 
capsule exserted from the often spreading 
calyx by 1/3-1/2 its length; calyx sparingly 
short-pubescent to glabrous. ... ...... ... 
Dee eee eee eee ees 6. L. deckertii Small. 

6. Aerial stems annual, herbaceous, dying to the 
base each year; capsule equaling the calyx or 
exserted not more than 1/5 its length from 
the closely enveloping sepals; calyx moder- 
ately to densely pilose. ............. sus. 7. 
7. Leaves abruptly tapering at apex into a 

hardened, shiny, conical callosity about 
0.25 mm. long; inner sepal clearly 3-nerved 
(often best demonstrated by moistening) ; 
pedicels averaging over 1.5 mm. long; 
capsule exceeding the sepals by about 1/5 
its length; seeds mostly 2. ... 7. L. pul- 
chella Raf. (= L. leggettii Britt. & Holl.). 
7. Leaves pointed but not differentiated into 
a callosity; inner sepal 1-nerved; pedicels 
averaging less than 1.5 mm. long; capsule 
almost completely enveloped by the sepals; 
seeds mostly 3-6. ...............00055 8. 
8. Calyces densely  appressed  pilose; 
stems and under surface of the leaves 

1974] Genus Lechea — Wilbur 481 

or at least the midvein and often the 
margins appressed pilose; capsules 
equaling or slightly shorter than the 
closely enveloping inner sepals. ...... 
PW ga EE 8. L. torreyi Legg. ex Britt. 

8. Calyces glabrous; stems and leaves 
completely glabrous; capsules very 
slightly exceeding the inner sepals at 
Maturity l.c vs vs 9. L. lakelae. 

Lechea lakelae Wilbur, sp, nov. 

Caudex simplex vel paucis aliquotve ramis; folia ramealia 
peranguste linearia-elliptica, acuta, 0.6-1.2 cm. longa et 
0.3-1.0 mm. lata, subtus costa et margine glabra; pedicelli 
glabri, (0.8) 1.0(1.2) mm. longi; calyx fructifer obovoideus, 
1.5-1.9 mm. longus et 1.2-1.6 mm. latus; sepala interiora 
glabra et leviter carinata, 1.5-1.9 mm. longa et 1.2-1.6 mm. 
lata; sepala exteriora glabra linearia vel anguste lanceolata 
1/2 vel 2/3 plo breviora quam sepala interiora; capsula 
c. 1.2-1.6 mm. longa; semina 3, c. 0.8 mm. longa. 

Caudex simple or with few to several branches; basal 
resting shoots unknown; aerial (i.e. flowering and fruiting) 
stems 1 to several, 2-3 (4) dm. tall, branching mostly above 
the middle and forming a rather compact, wiry-stemmed, 
bushy top with completely glabrous axes; cauline leaves 
lacking on all specimens seen; rameal leaves very narrowly 
linear-elliptic, tapering to both the apex and base, 0.6-1.2 
em. long and 0.3-1 mm. wide, completely glabrous above and 
below with the midvein somewhat elevated beneath; fruit- 
ing branches and pedicels glabrous, the pedicels (0.8)1.0 
(1.2) mm. long; fruiting calyx obovoid, broadest above the 
middle and tapering into the broadly obpyramidal, coria- 
ceous base, drying dark reddish brown, completely glabrous; 
inner sepals about 1.5-1.9 mm. long, including the indurate, 
obpyramidal base, and 1.2-1.6 mm. broad, slightly exceeded 
by the mature capsule, very broadly elliptic to almost or- 
bicular, apically broadly rounded, only the strongly elevated 

482 Rhodora [Vol. 76 

or slightly keeled midvein clearly visible; outer sepals linear 
to narrowly lanceolate, about 1/2-2/3 as long as the inner; 
capsules broadly barrel-shaped, cylindrical, about 1.2-1.6 
mm. long and 1-1.2 mm. in diameter, the valves firm, in- 
durate, tardily dehiscing about 1/2-2/3 the distance from 
apex to base; seeds 3, equilateral, dorsiventrally compressed, 
about 0.8 mm. long and 0.4 mm. wide, blackish. 

TYPE: Collier Co., Florida: Marco Island, moist level 
grassy area, higher beach of recently made lake. 7 August 
1967. Lakela 30953 (Holotype, DUKE; Isotype, USF). 

OTHER SPECIMENS EXAMINED: Collier Co.: coastal strand, Serenoa- 
Ceratiola scrub, growing with Cyperus in white sand, Lakela 31879 
(DUKE); Marco Island, coastal strand — Pinus elliottii association, 
Lakela & Almeda 31567 (DUKE); Marco Island with Indigofera, Poly- 
gonella, in Ceratiola-Quercus scrub, on U.S. 92, Lakela 27852A 
(usF); lagoon head, upper beach, Lakela 31673 (DUKE). 

My recollection and rather brief notes made more than a 
dozen years ago indicate that the species also occurs or 
occurred on Florida’s eastern coast in Broward Co.: sandy 
scrub above Fort Lauderdale, Buswell 24 July 1936 (Bus!). 

The apparent nearest relative of the new species, Lechea 
lakelae, is L. torreyi. The most obviously distinctive fea- 
tures are indicated in the key, but the two taxa may be 
readily distinguished at a glance. Obviously I feel that both 
merit recognition at specific rank. In the mind of some 
there still may remain a nagging suspicion that the striking- 
ly distinctive plants of the newly described taxon are merely 
a glabrous form of L. torreyi. I feel, however, that the ap- 
parently consistently longer capsules when compared to the 
length of the inner sepals suggest that L. lakelae is more 
than that. 

This species is named in recognition of Dr. Olga Lakela 
upon whose keen eye and careful collections our present 
knowledge of this taxon is largely based. It is humbling to 
recall that her recent significant contributions to south- 
eastern botany were made in her “retirement” years after 
an active career as a teacher and researcher in Minnesota. 

1974] Genus Lechea — Wilbur 483 

Not many botanists are destined to write floras of as distant 
regions as one of Minnesota's northeastern counties and 
Florida's southern tip. 

DURHAM, N.C. 27706 

WISCONSIN: This species, adventive from Eurasia, has 
heretofore been reported to extend from Newfoundland and 
New Brunswick to Delaware and the District of Columbia, 
west to Pennsylvania, Ohio, and Indiana (Mathias & Con- 
stance, 1944. Umbelliferae, in North American Flora 28B 
(1): p. 133; Fernald, Gray’s Manual, 1950; Gleason, Illus- 
trated Flora, 1952; Gleason & Cronquist, Manual of Vascu- 
lar Plants, 1963). Jones (Flora of Illinois, 1950) and 
Steyermark (Flora of Missouri, 1963) do not mention the 
plant, so it appears not to have spread westward. I report 
here its discovery in Wisconsin: WINNEBAGO CO.: roadside 
at junc. Country Club Road and Black Wolf Point Road, 
sect. 24, TI7N, R16E, 24 September 1967, Below 146; in 
roadside ditch ca. 144 mile from the junc. of county trunks 
YY and N, on N, sect. 20, T17N, R16E, 26 September 1972, 
Rhyner 044. 

The two specimens are deposited in the herbarium here 
at Oshkosh. The curators of the herbaria at MIL, UWM, 
and WIS have no records of the plant from Wisconsin; Iltis 
at WIS has kindly confirmed the identifications. The plant's 
occurrence at two sites four miles apart and spanning a 
five-year period warrants its inclusion as an element in the 
Wisconsin flora. 






Gametophytes of Lycopodium inundatum L., first re- 
corded for North America in 1972 (Bruce, 1972) from 
two localities in Michigan, were found growing at the edges 
of shallow wet depressions along the margins of a sandy 
access road on the shore of an inactive and flooded sandpit 
(Figs. 1, 2) in Somersworth, Strafford Co., New Hampshire. 
The mature sporophytes of L. inundatum var. inundatum 
(compare Gillespie, 1962) are abundant along the shoreline, 
often forming dense mats (Fig. 3). 

Samples of sandy soil containing the smallest visible 
sporophytes were collected in October and examined accord- 
ing to Bruce’s technique. Some of the young sporophytes 
were still attached to living gametophytes, as adjudged by 
the green color, turgidity, and characteristic lobing of the 
latter. Gametophytes were also found which had not yet 
produced sporophytes. 

Gametophytes ranged in size from less than one mm. in 
diameter for one which bore no sporophyte, up to about 
three mm. in diameter among those bearing sporophytes. 
Included in the latter category were one gametophyte bear- 
ing a sporophyte which had produced about ten microphylls, 
and two gametophytes each bearing two young sporophytes 
(Fig. 4). In contrast, however, some very young sporo- 
phytes were found to have no attached gametophytes, the 
latter apparently having rotted away, even though the 
sporophytes bore only one or two microphylls. This condi- 
tion appeared particularly prevalent where the soil particles 
were overgrown with moss protonemata and algae. 

In addition to variation in size discussed by Bruce, vari- 
ations in occurrence and condition, as well as in form (Bold, 
1973), of the gametophyte in relation to differences in the 

‘Published with the approval of the Director of the University of 
New Hampshire Agricultural Experiment Station as Scientific Con- 
tribution No. 756. 


1974] Lycopodium inundatum — Lane & Bogle 485 

microhabitat suggest the need for microecological studies 
in the field and laboratory. These would complement lab- 
oratory study of the culture and development of gameto- 
phytes of other species (Freeberg, 1962; Freeberg and Wet- 
more, 1957) and of the embryogeny of the sporophyte of 
Lycopodium inundatum (Bruce, 1972). 

The fact that spores of Lycopodium inundatum germinate 
readily and produce green photosynthetic gametophytes 
which live on the surface of the soil is of potential value to 
biology teachers (Bierhorst, 1964) in areas where the spe- 
cies occurs in New England (Fig. 5; for cireumpolar range 
map see Hultén, 1968, p. 26), for the gametophytes might 
be grown on artificially denuded patches of soil next to the 
sporophytes. Although gametophytes of mosses and ferns 
are readily available and commonly used for study in biology 
classes, few students ever see the live, fleshy gametophytes 
of Lycopodium. It is possible that careful searching in suit- 
able locations near colonies of mature sporophytes would 
produce numerous live gametophytes for class use. 

BrERHORST, D. W. 1964. Suggestions and Comments on Teaching 
Materials of the Non-Seed Bearing Plants. In: Vestal, P. A. 
et al. 1964. The use of living material in the teaching of botany. 
Amer. Biol. Teacher. 26: 89-113. 
BorLp, H. C. 1973. Morphology of Plants, Ed. 3. Harper & Row, 
New York. 688 p. 
BRUCE, J. G. 1972. Observations on the Occurrence of the Pro- 
thallia of Lycopodium inundatum. Amer. Fern Jour. 62: 82-87. 
FREEBERG, J. A. 1962. Lycopodium prothalli and their endophytic 
fungi as studied in vitro. Amer. Jour. Bot. 49: 530-535. 
, and R. H. WETMORE. 1958 (dated 1957). Gametophytes 
of Lycopodium as grown in vitro. Phytomorphology 7: 204-217. 
GILLESPIE, J. P. 1962. A Theory of Relationships in the Lyco- 
podium inundatum Complex. Amer. Fern Jour. 52: 19-26. 
HuLTÉN, E. 1968. Flora of Alaska and Neighboring Territories. 
Stanford Univ. Press. 1008 p. 
Seymour, F. C. 1969. Flora of New England. Charles E. Tuttle 
Co., Rutland, Vermont. 596 p. 


486 Rhodora [Vol. 76 

Figure 1. Typical macrohabitat with abundant mature sporo- 
phytes along brook (to left) and pond (to right) of access road 
(center). Note area of Fig. 2 (box). 

Figure 2. Microhabitat of gametophytes at edges of depression 
(left of center) darkened by growth of moss protonemata and algae. 

1974] Lycopodium inundatum — Lane & Bogle 487 


Figure 3. Vertical view of dense mat of mature sporophytes (each 
branch less than one cm. thick). 

Figure 4. Gametophyte bearing two young sporophytes (upper 
right dark area was green and minutely lobed, the remainder of the 
gametophyte was colorless). 

488 Rhodora [Vol. 76 

Figure 5. County distribution of Lycopodium inundatum varieties 
in New England (plotted from Seymour, 1969): var. inundatum 
(dots), var. Bigelovii Tuckerm. (circles), and var. robustum R. J. 
Eat. (triangles). Note site of gametophytes (X) discussed here. 



As a result of field and herbarium studies, a number of 
new or interesting records of plants have been found for 
Texas, Oklahoma, Kansas, and Nebraska. All specimens 
cited, unless indicated otherwise, are located at KANU. 

Corallorhiza wisteriana Conrad. Nebraska: DAWES CO.: 
Chadron State Park, open ponderosa pine wooded hillside, 
pine needle litter, clay loam soil, two stems growing in 
ravine, in fruit, 14 July 1972, Magrath & Weedon 7620 
(KANU). This collection represents an extension of the 
range of the coralroot southward from the Black Hills 
of South Dakota into the Pine Ridge of northwestern 

Holosteum umbellatum L. Texas: COLLINGSWORTH CO.: 0.3 
mi E of the Salt Fork of the Red River on T 203, growing 
in sandy soil in roadside ditch, scattered and uncommon, 
10 April 1973, Magrath & Richardson 7936 (KANU). KEN- 
EDY CO.: 18.3 mi S of Armstrong on US 77, growing in 
railroad right-of-way, 20 March 1972, Magrath, Rice & 
Johnston 7225 (KANU, OKLA). This collection represents 
an extension of the range of the species westward from 
Oklahoma into the Texas Panhandle and southward from 
Oklahoma to the Gulf Coastal area of southern Texas. 

Scleranthus annuus L. Oklahoma: GRADY CO.: Oklahoma 
College of Liberal Arts Campus (17th and Grand Avenue) 
in Chickasha; growing in flower bed on E side of Austin 
Hall; sandy soil; rare; associated with Arenaria, Bromus 
and Holosteum, flowers greenish-white, apetalous; 24 May 
1973; Magrath 8032 (KANU). This collection represents an 
extension of the range of this species southwestward from 
Missouri and eastern Kansas. 


490 Rhodora [Vol. 76 

Ranunculus testiculatus Crantz. Nebraska: DAWES CO.: 
Chadron State Park, 8 mi S of Chadron on Hwy #385, in 
campground in waste areas in shade of deciduous trees, 
weedy areas dominated by Taraxacum officinale, sandy soil, 
abundant, 6 June 1972, Weedon & Gates 7011 (KANU); 
Doris Gates, 29 April 1970. The range of this species is 
now known to be extended eastward into northwestern 
Nebraska. The above collection indicates the probable 
establishment of the species in abundance in this area. 

Thlaspi perfoliatum L. Oklahoma: KAY CO.: rest area on 
I-35, 2.8 mi N of Blackwell-Medford Exit; clay loam soil, 
somewhat sandy; small area near moist drainage ditch; 
only one colony seen, apparently introduced; associated 
with Bromus, Holosteum and Rumex; 6 May 1973; Magrath 
7969 (OKL). This collection represents an extension of the 
range of the species southwestward from Missouri and 

Eryngium prostratum Nutt. var. prostratum. Kansas: 
CHEROKEE Co.: 1.5 mi S of Galena on K 26, small cut-over 
and disturbed marshy area in Ozarkian oak-hickory woods, 
gravelly soil, associated with Juncus and sedges, apparently 
rare — only one plant found, prostrate vine rooting at the 
nodes, flowers bluish-purple, 22 October 1972, Magrath 7855 
(KANU). This collection represents an extension of the 
range of the taxon westward from the eastern Ozarks of 
Missouri, and northward from southeastern Oklahoma. 

Lactuca saligna L. f. ruppiana (Wallr.) G. Beck. Oklahoma: 
OKLAHOMA C0.: ca. 3 mi N of the Jct of I-44 and I-35 on 
I-35, a few scattered plants growing in road-cut, sandy 
loam soil, 5 August 1972, Magrath 7700 (KANU, OKL, OKLA). 
This collection represents an extension of the range of this 
taxon southwestward from Missouri and Kansas. 



Shortly after entering the Grand Savanna of Venezuela 
on the northern side we left the road to examine a sandy 
area with scattered small shrubs. On the patches of white 
sand between was a Drosera so small that it could easily 
be covered by a quarter and with so short a peduncle that 
the one or two flowers appeared practically sessile. This 
new species was such a happy augury for the Smiths' intro- 
duction to the Guayana Highland that we have christened 
it as follows: 

Drosera felix Steyermark & Smith, sp. nov. Fig. 1. 

Plantae solitariae; caulibus curtissimis; foliis dense rosu- 
latis, patentibus; stipulis ad petiolum 1/3 adnatis, ad me- 
dium 3-laciniatis, membranaceis, 2.5-3 mm longis, basi 
vaginantibus, solidis, 1 mm latis, segmentis laciniatis an- 
guste triangulari-lanceolatis, longe attenuatis, 1.7-2.2 mm 
longis, 0.2-0.3 mm latis, glabris; petiolis 5-5.5 mm longis, 
0.7 mm crassis, supra minute papilloso-verruculosis et pilis 
patentibus paucicellularibus acicularibus laxissime vestitis, 
subtus pilis multis adpressis simplicibus acicularibus onus- 
tis; laminis suborbicularibus, apice late rotundatis, 3 mm 
longis, 2.5 mm latis, castaneo-purpureis, supra margine 
glandulis 2-4 stipitatis et centro glandulis subsessilibus 
vestitis, subtus pilis eis petioli similibus; pedunculo nullo 
vel haud ultra 2.5 mm longo, eglanduloso; bractea lineari- 
lanceolata, acuminata, 1.5 mm longa, 0.1 mm lata, ciliata; 
floribus 1-2; pedicello 1-5 mm longo, pilis adpresso-adscen- 
dentibus simplicibus et glandulis remotis sessilibus atro- 
purpureis vestito; hypanthio cupulato-subcampanulato, 0.8 
mm longo, 1.2 mm lato, pilis patentibus remotis vestito; 
sepalis 5, lanceolatis, 1.5-1.8 mm longis, sub medium 0.8 mm 
latis, e pilis laxis ascendentibus simplicibus paucicellularibus 
hirsutulis; petalis 5, patentibus, albis, obovatis, apice ro- 
tundatis, basi cuneatis, 2.5 mm longis, 1.5 mm latis; stami- 


492 Rhodora [Vol. 76 

Fig. 1, a-h, Drosera felix: a, habit; b, single leaf (upper side); 
c, single leaf (lower side); d, hair detached from petiole; e, calyx 
and pistil with pedicel; f, stamen; g, petal; h, seed, ventral and 
dorsal views from left to right. 

1974] Venezuela Drosera — Steyermark & Smith 493 

nibus 5, quam petalis brevioribus; filamentis 1 mm longis, 
glabris; antheris subhemisphaericis, 0.5 mm longis, 0.4-0.5 
mm latis; ovario subgloboso, 1 mm longo, 1 mm lato, glabro; 
stylis 3, ad basim bipartitis, ramis 1.7 mm longis, glabris. 
Venezuela: BOLÍVAR: Carretera El Dorado to Santa Elena 
de Uairen south of El Dorado, alt 1200-1400 m, 19 Feb 1972, 
Julian A. Steyermark, Lyman & Ruth Smith, G. C. K. & E. 
Dunsterville 105468 (holotype VEN, isotypes NY, US). 


WASHINGTON, D. C. 20560 


Manuscripts should be submitted in duplicate and should 
be double-spaced or preferably triple-spaced (not on cor- 
rasable bond), and a list of legends for figures and maps 
provided on a separate page. Footnotes should be used 
sparingly, as they are usually not necessary. Do not indi- 
cate the style of type through the use of capitals or under- 
scoring, particularly in the citations of specimens, except 
that the names of species and genera may be underlined to 
indicate italics in discussions. Specimen citations should 
be selected critically especially for common species of broad 
distribution. Systematic revisions and similar papers should 
be prepared in the format of “The Systematics and Ecology 
of Poison-Ivy and the Poison-Oaks,” W. T. Gillis, Rhodora 
73: 161-237, 370-443. 1971, particularly with reference to 
the indentation of keys and synonyms. Papers of a floristic 
nature should follow, as far as possible, the format of 
“Contribution to the Fungus Flora of Northeastern North 
America. V.,” H. E. Bigelow & M. E. Barr, Rhodora 71: 177- 
203. 1969. For bibliographic citations, a recommended list 
of standard journal abbreviations is given by L. Schwarten 
& H. W. Rickett, Bull. Torrey Bot. Club 85: 277-300, 1958. 

Volume 76, No. 807, including pages 315-495, was issued Nov. 15, 1974. 


CONTENTS: — continued 
Two New Color-forms from Southern Maine 

Theodore W. Wells .......... eren nnne ener 
Range Extensions and New Records for the Bahama Flora 

Steven R. Hill 1... eese eene tornato tne eo osten atteso ases e tenet otto 

A New Species of Lechea (Cistaceae) from Peninsular 
Robert L, Wilbur ccccccccccccsscccccsccssecssecesseesnseceeeseeseessseeenseneeeeaeees 
Pimpinella Saxifraga L. (Umbelliferae) in Wisconsin 
Neil A. Harriman ....... eere eene enne eet tnnt neenon seno eno 

A Second Report of the Prothallia of Lycopodium 
inundatum in North America 

David M. Lane and A. Linn Bogle ............ ee 

New and Interesting Plants from the Central Plains States 
Lawrence K. Magrath and Ronald R. Weedon .................. 

A New Drosera from Venezuela 
J. A. Steyermark and L. B. Smith ........ eee 

Instructions to Contributors ............ eene 









Conducted and published for the Club, by 
ALFRED LINN BOGLE, Editor-in-Chief 


Associate Editors 

Vol. 76 December, 1974 No. 808 


Phenology and Physiognomy of the Hydrophyte Community 
in Otsego Lake, N. Y. 

Willard N. Hammam eene nnne nennt nter 

Panicum ensifolium Baldw. — New for Mexico 
Monn EONIA a iedera a REED 

Significant New Distributional Records for the Genus 
Sphagnum in the Northeastern United States 
Richard E. Andrus 


Environmental Control of Needle Characteristics in Sub- 
alpine Black Spruce 

Brian F. Chabot 


(Continued on Inside Back Cover) 




The Nem England Botanical Club, Jne. 
Botanical Museum. Oxford St.. Cambridge. Mass. 02138 

RHODORA. — A quarterly journal of botany, devoted primarily to the 
flora of North America and floristically related areas. Price $20.00 
per year, net, postpaid, in funds payable at par in the United States 
currency at Boston, Some back volumes, and single copies are 
available. For information and prices write RHODORA at address 
given below. 

Scientific papers and notes, relating directly or indirectly to the 
plants of North America, will be considered by the editorial com- 
mittee for publication. Articles concerned with systematic botany 
and cytotaxonomy in their broader implications are equally accept- 
able. All manuscripts should be submitted in duplicate, and should 
be double-spaced throughout. Please conform to the style of recent 
issues of the journal. Illustrations can be used only if the cost of 
engraver's blocks is met through the author or his institution. 
Forms may be closed five weeks in advance of publication. Ex- 
tracted reprints, if ordered in advance, will be furnished at cost. 

Address manuscripts and proofs to 

Dept. of Botany, University of New Hampshire, 
Durham, New Hampshire 03824 
Subscriptions and orders for back issues (making all remittances 
payable to RHODORA) should be sent to RHODORA, Botanical 

Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive 

the next number of RHODORA, changes of address must be received 

prior to the first day of March, June, September or December. 
Second Class Postage Paid at Boston, Mass. 




Vol. 76 December, 1974 No. 808 



As part of a biological survey conducted in 1969, the 
species and distribution of aquatic macrophytes in Otsego 
Lake, Otsego Co., N.Y., were determined (Harman 1970). 
The 1969 information was compared with data collected by 
Muencher (1936) in 1935 (Harman & Doane 1970). Emer- 
gent plants were more abundant in 1969 than in 1935. The 
number of species present in the lake had decreased between 
1935 and 1969 and an introduced species, Potamogeton 
erispus L., that was not present in 1935, was the most 
obvious plant in the lake in early summer of 1969. The 
reduction in species in this lake is similar, although not as 
pronounced, as changes observed over 50 years in Oneida 
Lake, N.Y. (Harman & Forney 1970) and over 70 years in 
Put-in-bay Harbor, Lake Erie, Ohio (Stuckey 1971). 

During the summer of 1969 it was noted that the zooben- 
thos and zoo-meroplankton populations underwent drastic 
fluctuations that correlated with the phenology of the aqua- 
tic macrophytes and the seasonal changes in the physiog- 
nomy of the plant community. A study was initiated to 
more accurately determine the composition of the flora at 
various times during the year so that we could quantita- 
tively ascertain its effects on the associated fauna. The 
purpose of this report is to describe the seasonal changes in 


498 Rhodora [Vol. 76 

the appearance of the littoral zone in Otsego Lake at Rat 
Cove, at the SUNY Oneonta Biological Field Station at 
Cooperstown, New York. 


In 1970, a transect about 500 m long was plotted from 
the end of our docks through the littoral zone, to the deeper 
water beyond. Plants growing along this line were meas- 
ured by divers each week, from 1 June through 2 Septem- 
ber, to determine the emergence time of the spring cohorts, 
greatest maximum heights attained, times of flowering, and 
times of death and decomposition of each species (Harman 
1971). Because all growth represented even-aged stands, 
sufficient data were obtained by measuring the tallest in- 
dividual plants of each species each week. It was impossible 
to measure the same individuals throughout their existence 
because the act of measurement disrupted, if not destroyed 
these fragile organisms. Also, movements along the tran- 
sect disturbed the sediments so that by the end of Septem- 
ber we had to move farther and farther away from the 
original line in order to measure plants in an undisturbed 
environment. Secchi transparency and surface water tem- 
peratures in Rat Cove were recorded from 15 June through 
25 August. 

In 1971 the same procedures were followed with these 
changes and additions. Samples were taken through the 
ice on 5 February. Weekly observations were initiated 
Shortly after the ice breakup (22 April) and continued 
until 16 December after all plants had completed their 
seasonal growth. 'The transect was marked with an an- 
chored hand line for divers to follow in order to alleviate 
the problems of getting lost in the often